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O20 NDE, 

1907, pp. 1-446. 





Aces Sill 








His Grace THe Duke oF BeprorD, K.G., President. 

Grorce A. BouLencer, Esq., 
F.R.S., Vice-President. 

JOHN Rose Braprorp, Esq., 
WILDS; IDESCh, IeHSh. oes 

F. Dawrrey Drewirr, Esq., | 

M.A., M.D. 

CHARLES Drummond, 

Str Epwarp Duranp, Br., C.B. 

F. Du Cane Gopman, Ksq., 
D.C.L., F.R.S., Vice-Presi- 

M.A., F.R.S. 

Sir EpmunpD Gites Lover, Br., 

Pror. Epwarp ALFRED M1v- 
cuin, M.A. 



P. CHatmers MitcHey, Esq., 
MA. (DIS coins SHS 

W. R. Oaitvir-Grant, Esa. 

ALBERT Pam, Esq. 

E. Lort Puiwuirs, Esq. 

Sir Parrick Puiayratr, C.1.E. 

D.Sc., M.P. 

Howarp Saunpers, Hsq., Vice- 

Davin Seru-Smiru, Hsq. 

OLDFIELD THomas,  KEsq., 

A. Trevor-Battys, Esq., M.A. 

Henry Woopwarp, Ksq., LL.D., 
F.R.S., Vice-President. 

P. CHatmers Mircnrett, M.A., D.Sc., LL.D., F.BS., 


Frank E. Bepparp, M.A., F.R.S8., Prosector. 
R. I. Pocock, F.L.8., Superintendent of the Gardens. 


F. H. Waternouss, Librarian. 

JoHN Barrow, Accountant. 
W. H. Cots, Chief Clerk. 


RAC as 

GrorcGe ArtHur Dovusiepay, Clerk of Publications. 
ARTHUR THOMSON, Assistant Superintendent of the Gardens. 


1907, pp. 1-446. 

January 15, 1907. 

The Secretary. Report on the Additions to the Society’s 
Menagerie during the months of November and De- 
Gemlboer WO OG spas acer eyeysclck a cacleee re rar ege ce pice eens ame sms 

Mr. Oldfield Thomas, F.R.S., F.Z.S. Description of a new 
Monkey from the Ituri Forest. (Plate I.) ............... 

1. On a Collection of Mammals made by Dr. Vassal in 
Annam. By J. Lewis Bonnore, M.A., F.LS., F.Z8. 
(Bibi ire Ul) pears ARs SMR UB Oe ane ae aes enP One Aloe Sa Or Pose 03 

2. On the “ Bleating” or ‘“ Drumming” of the ee 
(Gallinago celestis). By P. H. Baur, B.A., F.ZS. . 

3. Contributions to the Knowledge of the Systematic 
Arrangement and Anatomy of certain Genera and 
Species of Squamata. By Frank E. Bepparp, M.A., 
BUR: Se eerosecsorsortive: SoOcietige.s-6. dae ace eteen eee 

4, A List of Moths of the Family Pyralide collected by 
A. E. Pratt in British New Guinea in 1902-3, with 
Descriptions of new Species. By Grorce H. Kenrick, 

F.Z.S. (Plates IIT. & TRV SaaS ten tine Senin ten ce b ies oa ob 
a 2 






5. On some new and insufficiently known Species of Mar- 
moset Monkeys from the Amazonian Region. By Prof. 
Dr. Emit A. Gorupt, C.M.Z.S., Director of the Para 

SO oe mee reese meee eee sees eseeeees es esaeeeesseseeresssersvsesee 

February 5, 1907. 

Mr. F. Martin Duncan. | Cinematograph exhibition of 
Animals in the Society's Gardens and other Zoological 
SIU OU I STCUS arlene ere tie) CRD CPP Ee AREA nA ShE let ac .509 

Mr. Oldfield Thomas, F.R.S., F.Z.S. Exhibition of a collec- 
tion of Mammals and Birds from the Islands of Saghalien 
and Hokkaido 

Cem ee mew twee mes wesc eee neers ease eee se ese ese es teserene 

Dr. W. T. Calman, F.Z.8. Notice of a paper on New or 
Rare Crustacea of the Order Cumacea from the Collection 
of the Copenhagen Museum 

1; The Origin of the Lateral Horns of the Giraffe in Fetal 
Life on the Area of the Parietal Bones. By E. Ray 
Lanxester, M.A., D.Sc., LL.D., F.R.S., F.Z.S., Director 
of the Natural History Departments of the British 

2. Parallel Hair-fringes and Colour-striping on the Face of 
Fetal and Adult Giraffes. By E. Ray Lanxsster, 
MA’, DSc., UE.D., EF.R.S:, F:Z.8., Director of the 
British Museum (Natural History). (Plate V.) 

3. On the Existence of Rudimentary Antlers in the Okapi. 
By E. Ray Lanxester, M.A., D.Sc., LL.D., F.R.S., F.Z.S., 
Director of the British Museum (Natural History). 
((Bbites VAI GG BY Wi th PP Ok He kine ah a aA 

4. Description of Hyla resinifictrix Goeldi, a new Amazonian 
Tree-Frog peculiar for its Breeding-habits. By Prof. 
Dr. Emmi A. Gortp1, C.M.Z.8., Director of the Para 

5. The Duke of Bedford’s Zoological Exploration in Eastern 
~, Asia,—IIT. On Mammals obtained by Mr. M. P. 
Anderson in the Philippine Islands. By Otprieip 
MroMAS, HORS. EeZis. 








February 19, 1907. | 

The Secretary. Report on the Additions to the Society’s 
Menagerie during the month of January 1907 

Dr. C. I. Forsyth Major, F.Z.8. Exhibition of remains of a 
Bear trounan Cay ernie Consicay nese crre eee enn 
1. On English Domestic Cats. By R. I. Pocock, F.LS., 
F.Z.8., Superintendent of the Zoological Society’s 
Gardens. (Plates VITI.—X.) 

eee et eee ree eee estes eee ses eseseee 

2. Report on Deaths occurring in the Society’s Menagerie 
during 1906. By C. G. Srniemann, M.D., F.ZS., 
Pathologist to the Society 

Pepe meee eee e enero reese reese sereeseese 

3. On a peculiarly Abnormal Specimen of Turbot. By J. T. 
CunnincHam, M.A., F.Z.S8. (Plate XI.) 

eee eee reece se esece 

4. On the Azygos Veins in the Mammalia. By Frank E. 
Bepparp, M.A. (Oxon.), F.R.S., Prosector to the Society 

5. Ideas on the Origin of Flight. By Dr. Baron Francis 

March 5, 1907. 

The Hon. Walter Rothschild, M.P., F.Z.8. Exhibition of a 
mountedyspecmmenior a) Govillaigetacac-acesa-i crise cee se 

1. Descriptions of a new Species and two new Subspecies of 
Antelopes and a new Sheep. By the Hon. Water 
RopHscHiED Mims (Phas sHI ZR Stee ge oiiy. ds ceeaeaectias setae 

2. On Elephant Remains from Crete, with Description of 
Hlephas creticus, sp. n. By Dorornna M. A. Bare. 
(Plates XII. & XIII.) 

eee eee etme etree reser eons ee ess sess ees assoee 

3. Zoological Results of the Third Tanganyika Expedition, 








conducted by Dr. W. A. Cunnington, 1904—1905.— » 

Report on the Polyzoa. By Cuarztes F. Rovusseer, 
FRLMS. (Plates @kViGiXVs)08 te. Ae aelee tenes 


4. Zoological Results of the Third Tanganyika Expedition, 
conducted by Dr. W. A. Cunnington, 1904-1905.— 
Report on the Brachyurous Crustacea. By Wrixiam A. 
Cunnineton, B.A., Ph.D., F.Z.S. (Plates XVI. & 

Ome mee w et eee eet eee reer reese een e te essere een eee ener ete seesu eee 

5. On Two new Species of the African Genus Microchetus 
belonging to the Collection of Oligocheta in the Museum 
of Christiania. By Frank EH. Bepparp, M.A., F.R.S., 
Prosector to the Society 

March 19, 1907. 

The Secretary. Report on the Additions to the Society’s 
Menagerie during the month of February 1907 ......... 
Mr. Herbert F. Standing. Notice of a Memoir on recently 
discovered Subfossil Prosimiz from Madagascar 

eee ee eeee 

1. Descriptions of some New Species of Animal Parasites. 
By L. W. Sampon, M.D., F.Z.8. 

2. Descriptions of five New Species of Hemogregarines from 
Snakes. By L. W. Sampon, M.D., F.Z.S., and C. G. 
Seviemann, M.D., F.Z.8. 

ii ii i i ii i i iii i aii ai ee ay 

3. The Rudd Exploration of South Africa—VII. List of 
Mammals obtained by Mr. Grant at Coguno, Inhambane. 
By Ouprietp Tuomas, F.R.S., F.Z.S., and R. C. 
Wrovuenton, F.Z.S. 

April 9, 1907. 

Mr. R. I. Pocock. Exhibition of a photograph and the skull 
of a specimen! of Pallas’s Cat that had recently died in 
the Menagerie, with general remarks on the species 

1. On a small Collection of Fishes made in the Eastern 
Watershed of the Transvaal by Capt. G. E. Bruce. By 
G. A. Bounencrr, F.R.S., F.Z.S. (Plates XVIII. & 

2. On the Winter Habits of the Greater Horseshoe, Rhino- 
lophus ferrum-equinwm (Schreber), and other Cave- 
haunting Bats. By T. A. Cowarp, F,Z.8. ............... 








-. 2M) 


3. Notes upon the Anatomy of aSpecies of Frog of the Genus 
Megalophrys, with references to other Genera of 
Batrachia. By Frank E. Bepparp, M.A., F.RS., 
Erosector toute; Society mn-reneeceeeeceai-sndassocse ae 324 

4. Contributions to the Osteology of Birds.—Part IX. 
Tyranni; Hirundines; Muscicape, Lani, and G'ymno- 
nhines., ByaNVis Be Py Crani kh Anse eAelunS OCC mesa -ee ae 352 

April 23, 1907. 

The Secretary. Report on the Additions to the Society’s 
Menagerie during the month of March 1907 ............ 319 

Dr. A. Smith Woodward, F.R.S., F.Z.S. Exhibition of a 
malformed antler of the Red Deer ....................000000% 380 

Mr. R. I. Pocock. Exhibition of a model of the African 
J Dlg lave at Go 0 Xs dee RP ARE irri, ocs ooceb accsucdscmoretoe’ 380 

Mr. C. J. With. Notice of a Memoir entitled “* An Account 
of the South-American Cheliferine in the Collections of 
the British and Copenhagen Museums” .................. 380 

1. The Ears as a Race-Character in the African Elephant. 
BR Sb yD ORE ie said nis ain ie Sager n ete acne stele aeieotenbercine mses 380 

2. The Duke of Bedford’s Zoological Exploration in Eastern 
Asia.—IV. List of small Mammals from the Islands of 
Saghalien and Hokkaido. By OuprreLtp Tuomas, F.R.S., 
F.Z.S. (With Appendix on the Cold-blooded Verte- 
brates, by G. A. Bounencer, F.R.S., F.Z.8.)............... 404 

3. On some new Species of Earthworms of the Family 
Kudrilide, belonging to the Genera Polytoreutus, Neu- 
mannielia, and Hminoscolex, from Mt. Ruwenzori. By 
RANKS Hee BEDDARDs UV AY. THES user io: asee sda seeene aces A415 

4. Some Notes on Hybrid Bears. By Henry SCHERREN, 
BWA cgrtae reno s acide ices notte Na tlea har vst sls Vaccines snaee eee 431 





With References to the several Articles contributed by each. 

1907, pp. 1-446. 

Baur, Purip Hernricn, B.A., F.Z.8. 
On the “Bleating” or “ Drumming” of the Snipe 
(Gallinage cochestis) .....0......ccsceuecee neces nen ene eenneceentees 12 
Bare, Miss Dororuea M. A. 
On Elephant Remains from Crete, with Description of 
Elephas creticus, sp. n. (Plates XII. & XIII.) ...........: 238 
Bepparp, Frank E., M.A., F.R.S., Prosector to the Society. 
Contributions to the Knowledge of the Systematic 
Arrangement and Anatomy of certain Genera and Species 
Fok. Sofuaniarbaly eerste siesta os Leet vealed 30 
On the Azygos Veins in the Mammalia ............-..++- 181 

On Two new Species of the African Genus Microchetus 
belonging to the Collection of Oligochzta in the Museum 
Of Christiania. «acaeaesedsdesapoedsanseee Sasdacetemeesesen gene ee ee. SET 

Bepparp, Frank E. (Continued.) 

Notes upon the Anatomy of a Species of Frog of the 
Genus Megalophrys, with references to other Genera of 
IBEW A GX 0) 00 nar neat RMR ear MOE ator ons sAhdsea sec odoox 

On some new Species of Earthworms of the Family 
Hudrilide, belonging to the Genera Polytoreutus, New- 
manniella, and Hminoscolex, from Mt. Ruwenzori 

Bonuote, J. Lewis, M.A., F.LS., F.Z.8. 

On a Collection of Mammals made by Dr. Vassal in 
Aeninrermmy (@Plaibe ET, \? oo Seca) een: Seren eee Ut). 9 

Bou.encer, Grorce AuBert, F.R.S., V.P.Z.S. 

On a small Collection of Fishes made in the Eastern 
Watershed of the Transvaal by Capt. G. E. Bruce. 
(Plates XVIII. & XIX.) 

Bee www eee eet were re eee e er esse se en sesene 

On the Cold-blooded Vertebrates of the Islands of 
Saghalien and Hokkaido. See THomas, OLDFIELD. 

Catman, WittiAm Tuomas, D.Sc., F.Z.S., of the British 
Museum (Natural History). 

Notice of a paper on New or Rare Crustacea of the 
Order Cumacea from the Collection of the Copenhagen 
Museum: e222 eek) Re RY a ey I 

CowarpD, THomas ALFRED, F.Z.S. 

On the Winter Habits of the Greater Horseshoe, 
Lhinolophus ferrum-equinum (Schreber), and other Cave- 
hauintns Bats) 4h sees ckicclsncus dst aT ee ee 

CUNNINGHAM, JosepH THomas, M.A., F.Z.S. 

On a peculiarly Abnormal Specimen of Turbot. 
(Plate XCM), 3:3... 385 das fede eae = gee ee 







Zoological Results of the Third Tanganyika Expedi- 
tion, conducted by Dr. W. A. Cunnington, 1904-1905. 
—Report on the Brachyurous Crustacea. (Plates XVI. 
gC 2 I) es ie Re coro cn tL A oe ee 

Duncan, F. Martin. 

Cinematograph exhibition of Animals in the Society’s 
Gardens and other Zoological Subjects 

GoEtp1, Prof. Dr. Emi A., C.M.ZS. 

On some new and insufficiently known Species of Mar- 
moset Monkeys from the Amazonian Region 

Description of Hyla resinifictria Goeldi, a new Ama- 
zonian 'Tree-Frog peculiar for its Breeding-habits 

Kenrick, Grorce Hamiuron, F.Z.S. 

A List of Moths of the Family Pyralide collected by 
A. EK. Pratt in British New Guinea in 1902-3, with 
Descriptions of new Species. (Plates III. & IV.)......... 

Lankssrer, Sir Epwin Ray, K.C.B., M.A., D.Sc., LL.D., 
F.R.S., Director of the British Museum (Natural 

The Origin of the Lateral Horns of the Giraffe in 
Foetal Life on the Area of the Parietal Bones ............ 

Parallel Hair-fringes and Colour-striping on the Face 
of Fetal and Adult Giraffes. (Plate V.) ...............08. 

On the Existence of Rudimentary Antlers in the Okapi. 
(WHEE a iaiwin WAUL5) gaoesbuccedesbeccooocba64 oogucosboaedddiconde 








Lyprexxer, Ricuarp, B.A., F.R.S., F.Z.S. 

The Ears as a Race-Character in the African Elephant 

Magor, Dr. C. I. Forsyru, F.Z.S. 

Exhibition of remains of a Bear from a Cavern in 

Si) =heiesele.2)\2/.eie)s) aces ie ie)alaralei=)\sieleleleiele/e)\ele\s!a 0/6 o/slelle/«ielelalelsisisselalalelereienerehciniatctate 

MircHEtt, Peter CHALMERS, M.A., DSe., LD. F.R:S., 
Secretary to the Society. 

Report on the Additions to the Society’s Menagerie 
during the months of November and December, 1906 


Report on the Additions to the Society’s Menagerie 
during the month of January 1907 

Cece er cece reese esse secs eces 

Report on the Additions to the Society’s Menagerie 
during the month of February 1907 

Cece rsescesesvecesscssvesece 

Report on the Additions to the Society’s Menagerie 
during the month of March 1907 

Seer cee errors so recess esses cene 

Norcsa, Dr. Baron Francis. 

Ideas on the Origin of Flight 

Set eee ee sere secsccccre esse nsec 

Pocock, Reervaup Iynzs, F.L.S., F.Z.8., Superintendent. of 
the Gardens. 

On English Domestic Cats. (Plates VIII.-X.)......... 

Exhibition of a photograph and the skull of a specimen 
of Pallas’s Cat that had recently died in the Menagerie, 
with general remarks on the species 

ers eeeeercccoereesaovess 

Exhibition of a model of the African Elephant 
“« Jumbo ” 

see ee oes e ses cescceresesesessovrersseesoceresocssceeesse essences 










Pycrarr, WitLIAM Puaneg, F.Z.8., A.L.S8., of the British 
Museum (Natural History). 

Contributions to the Osteology of Birds.—Part IX. 
Tyrannt; Hirundines; Muscicape, Lanti, and Gymno- 

Roruscuitp, The Hon. L. Watrer, M.P., D.Sc., Ph.D., 

Exhibition of a mounted specimen of a Gorilla ...... 237 

Descriptions of a new Species and two new Subspecies 
Ot AMIE DES cinl Er SINEEO ooncoossdesosceonnuosoacss0e 6 Sead 


Zoological Results of the Third Tanganyika Expedition, 
conducted by Dr. W. A. Cunnington, 1904—1905,— Report 
Gin Woe) IHolbywor,. (Ede NsGiss SUING ay DW ons easodoereenace: 250 

Sampon, L. W., M.D., F.ZS. 

Descriptions of some New Species of Animal Parasites. 282 

Sampon, L. W., M.D., F.Z.8., and Seuiegmann, C. G., M.D., 
E.Z.8., Pathologist to the Society. 

Descriptions of five New Species of Hemogregarines 
EPROM SMA ROS Me eee ste arlene hole ieee eee lata se sas dala hela ae aun Sa: 283 

ScHERREN, Henry, F.Z.S8. 

Some sNoteston aly bridmbearcm eae era eee ee ee 431 

Seviemann, C. G., M.D., F.Z.8., Pathologist to the Society. 

Report on Deaths occurring in the Society’s Menagerie 
Glpiaine? IMG ope seadessnnesdoasbenssbenouadde cose | codudasdnscssnnoc0s 

Seriemann, C. G., M.D,, F.Z.8., Pathologist to the Society, 
and Samson, L. W., M.D., F.Z.S. 

Descriptions of five New Species of Hemogregarines 

from Snakes..... Sr ROE NS IE la crc SO RE 3 

Sranpine, Herpert F., M.Sc. 

Notice of a Memoir on recently discovered Subfossil 

TPrpogiraes tort INENORYSEING A 5 occccebacucco does apcocs00n030006 

Tuomas, OLDFIELD, F.R.S., F.Z.S8. 

Description of a new Monkey from the Ituri Forest. 
(Plate VL.) 2.........ce0 seen ee cee tee tee ee ccee eters cents ectesecaeeeners 

Exhibition of a collection of Mammals and Birds from 

the Islands of Saghalien and Hokkaido ..................... 

The Duke of Bedford’s Zoological Exploration in 
Eastern Asia.—III. On Mammals obtained by Mr. M. 
P. Anderson in the Philippine Islands7..5..2.......0.......-- 

The Duke of Bedford’s Zoological Exploration in 
Eastern Asia. —IV. List of small Mammals from the 
Islands of Saghalien and Hokkaido. (With Appendix on 
the Cold-blooded Vertebrates, by G. A. Bov.encrr, 
TRISH AISE)) Scone vengspeusaaonocodsc0:-. he aise 

‘Tomas, OLDFIELD, F.R.S., F.Z.S., and Wrovueuton, R. C., 

The Rudd Exploration of South Africa,—VIT. List of 
Mammals obtained by Mr. Grant at Coguno, Inhambane. 










Wirn, C. J., of Copenhagen. 

Notice of a Memoir entitled “*An Account of the 
South-American Cheliferine in the Collections of the 
British and Copenhagen Museums tl a, se Ree ee 

Woopwarp, Artuur Suiru, F.R.S., F.Z.S., Keeper of the 
Geological Department of the British Museum. 

Exhibition of a malformed antler of the Red Deer 


Wrovueuton, R. C., F.Z.S., and Tuomas, OLDFIELD, F.R.S., 

The Rudd Exploration of South Africa—VII. List of 
Mammals obtained by Mr. Grant at Coguno. Inhambane. 




TI. | 
IOV {| 





190i pps Elo: 

Cercopithecus dentt ecu ce esse cc essere ete 2 
NycEicebUs PYGMEUS 2.256. 0 cece eset et te ees 3 
Pyralidee from British New Guinea .........+++..0-+> 68 
Head of a Fetal Giraffe (half the natural size) ......-- 115 
torn=tipsy orm Okerpietrs eet iarier so co vsatolee: (198 
Young Ossicone or Horn of Okapi..........-+. 200000 Fi 
Blotched Tabby, Felis catus ......--..e eset ] 
Striped Tabby, Felis torquata ..........00 sees beeen , 143 

Agriotypes of the Striped Tabby 

Nbnormal VouneMurbot 2: 4. oti et 174 
Elephant Remains from Crete........00.+.s.-eee eres 238 
Tanganyika Polyzoa ........--.0-ee eee eer s teens 250 

1. Potamen (Potamonautes) orbitospinus. 2-7. Antennal 
Region of various Potamonidee .......-..--+++++> A 

1&3. Potamon (Potumonautes) platynotus. 2 & 4. Platy- | a 
thelphusa conculeata. 5 & 6. Platythelphusa maculata 

1. Barbus bructt. 2. B. sector... 00.0. cee e nese sence | « 

Varicorhinus brucit . 6.0... eect ees: \ 

Foou. Sov.— 1907. h 

Vial) ibn cca 
ae: en ‘lib Res dept sgn 

yeti! Mirksiies 


ae, Bi oe ie 

re Ey atten 

eect pest 
| “hh pdt A 6 oO vabinalatiagl 
ff. ee Cee sn a 

Das) ot Mee ee 

C2 bo 


1907, pp. 1-446. 

. Palatal view of skull of Nycticebus pygmeus.. cc... ceveen. 
. Lateral view of skull of Mycticebus pygmeus... 0... cece eee 
. a. Snipe bleating, showing characteristic position. 6, Formation 

Oli tallbaszordimeanilw ibe! deinetic: hime re teenie 

. Varieties of outer tail-feathers of Gallinago cwlestis ........ 
. a, Dorsal view of musculature of tail of Snipe. 6. Ventral 

view of musculature of tail of Snipe ...............00008 

. A. Half of tail of Gallinago celestis from without inwards 

left to right. B. Half of tail of G. delicata from without 
inwards left to right. C. Half of tail of G. gullinula ...... 

. A. Section of two rami of a feather showing interlocking of 

distal and proximal radii. B. Ramus of Gallinago celestis, 
showing proximal and distal rows of radii. C. Distal radius 
of G. celestis. D. Proximal radius. E, Distal radius of 

middle-tail feathers of G. calestis.........0. cc eves seeeees 

8. A. Half of tail of Gailinge major from without inwards left 
to right. B. Half of tail of G. nobilis from without inwards 

left to might ©: Elalfion tailot Ge stenw@ 55.0405 

9, a. Tail of Gallinago solitaria. 6. Tail of G. megala ........ 
10. Lung of Chameleon verrucosus, entire ........000 0. eevee 
ie Wune ot Chameleonidilepis entire wens. + ssc ls. eee 
12. Lung of Chameleon parvilobus, opened longitudinally ........ 
13. Lung of Chameleolis, opened longitudinally ................ 
14, A. Stomach and pancreas of Chameleolis. B. Dorsal aorta 
ATG MAMIE Deine ENE No LE BWasbaeoeseoedul si cow we awe 
l5eMkgua sconcowdes Weadand meck sai j4. 2 oes 
16, A portion of the trachea, the bronchi, and the upper parts of 

the lungs of Heloderma, fvom the dorsal side .............. 







be te woe el 
= S 

bo bo 
Co bs 











. Pancreas, gall-bladder, &c., of Varanus niloticus, to show the 
course of the bile-ducts .......... a haa pial ele Sata ea GI tice 64 
IDO) LE [Arie WIS CMO OTOCHOHIES. ooo co ne@adounsoobannebasor 65 
PUG SOL WAKAWUS GIUSEUS). .. «5. o)eeMeenne anon oe eae eae 66 
Mbackeviewsothead Olu VVd as C/(OUtes ime aetna 89 
Mehack views of headuot Midas 7 /Ucentcrassmr ere eee 6 Gil 
A Backvie, of lead of Mudasigniscoventea 1. eee seer 93 
milileadsote Midas nye ator |. sateen 94 

. Lateral view of the skull and lower jaw of a very youny 

Giralle. sis wane ero anes eee © Pongtie  aeep aan 101 

, Lateral view of the skull of a Giraffe, about two-thirds grown. 102 
26. View from above of the fronto-parietal region of the skull of a 

WEIAP ROUTES EDN RNIE Rea gaoo suo dando ds cl Gon OO HdMmaD eo bG oA. 102 

. View from above of the fronto-parietal region of the skull of an 

duane tute MO Ka pi wis aveieraagueva tse, asa tor occ usteenemetewerstate terete suelo seecett 103 

28. Sagittal section through the bony tissue of the ossicone and 

the roof of the skull of a very young Giratfe .............. 104 

. Copy of the drawing of a sagittal section through the ossicone 

and the roof of the skull of a newly-born Giraffe, published 
by the late Sir Richard Owen in the Trans. Zool. Soc. 1840.. 104 

. Drawing of the right side of the head of the fcetal Giraffe 

described in the text .......:....... BASES. 6 SOR wae 106 

Uhesleic shormesoiethe) tcetalmGntalcmnn tyne irre eo nian ot 107 
. Diagram to show the flattened plate-like form smi the 

orientation of the horns ef the foetal Giraffe .............. 108 

33. Skull of Giraffe....showing the directions in which the 

Slut welsh CULM: fraatre fea Deocaerensrn catnenewes cisia jes Rerpiere Meets 110 

. View, looking forwards, of the skull cut in the direction of 

ivory 424 a WORE SECM SE CMbiclke asa wa occ onesies codahblds Dak dil 

. Drawing, of the natural size, of a young foetal skull of a 

(CHIPTEHE Reta ae eS eR amen ae aes aR ake cha Bis Hines ers Se ey nae 112 
Diagram to show the various positions on the frontal bone at 
which the bony horn-cores of the Cavicorn and Cervine 
Tuminants may take them erowth\..- s-)) e eee reece - 114 
Left side of the head of the foetal Giraffe described, showing 
colour-stripes on the snout and above and below the eye.... 115 

. Section across three dark-coloured bands above the eye of the 

foetal Giraffe, showing three longitudinal furrows or de- 
pressions in transverse section corresponding to the dark 

WHC mc oeadescetesero vos cs ado essen seovsdbecosagmaaE 116 
Section of the skin of the frontal region of the foetal Giraffe, 
showing threesizes Of Waits Terr miele elaine ela aie-/ ( Lily 

Surface-view of banded “pelage”’ of the frontal region of foetal 
Giraffe, showing the convergence of the hairs at the three 
longitudinal Tiana Oi Gawd ayyOCRMNNOS 5 ocdococsod.essccnce 118 

An enlarged drawing of a single hair of average size ofr om the 
frontal region of an adult Giraffe, showing the ‘dark pigmented 
free extremity and the opaque white lower DOAN “S sueuees 119 


; Colour-striping on the face of a Somali-land Beant . chain teegiiths 124 
. The same specimen as that shown in text-fig. 47, with colour 

Colour-striping on the face of an adult male Giraffe from 
Kordofanvirraa i) 4. ae ey Sepak st, Atl areecare Hiaiith 120 

. Colour-striping on the face of an adult female Giraffe from 

ond ofan ree Oy Soy Pay hou edo ceaneapebisieh aisha bis 120 

. Colour-striping on the Anas al Giraffa ae dalis cottont 

from: Mitz HleonsUWeanday icy anne ‘ RU avant ata Ay 

: Colour-striping on th face and muzzle ofa a Thaiernsel Giraffe... 122 
. The same specimen as that shown in text-fig. 45, with colour 

OMMUETERE: Aye Ye). CON aeaee Bvt capay betas Fe Reet ee ile 

Onna GS Pea ol I SRS. 8 Nk CRY EN) ecu NURS ea ie HERR 124 

. Drawing of a fore and aft section through the tip of the ossicone 

of amt adults Okapi | Ase A) eects: & skaters byevenateverenne as tasers acer 126 

5), Rudimentary free ossicone of hemispherical shape from the 
skin overlying the frontal bossed region of the skull of an 
Okapitofethe broud-siculleditype) Wank aes see aces niet 128 

51. Section of the ossicusp drawn in text-fig. 50, to show the 

In{coOmlpleter ossilie sti Om mej vecvalss 9 -)-Naate ene enekeielsy siotaet ate cist etele 128 

52. Bisected horn-tip of the Edinburgh Okapi...............055 13L 

53. Section of half of the same ‘Waecal OMM=bip Ns. 1 « 131 

64. Drawing of the right ossicone of the Okapi brought by Capt. 
Boyd Alexander “Fhonin the Welle River ...... Maint: tovetoers 154 

55. Section through the tip of the same specimen to show the 
incomplete trabeculated Ossification’............---.s++5+- 134 

DGwerylal resinafict) Uy Mule wit acts ei qumGics shiseier ni eke a aye 136 

is Jalil Tay vonee (Cheerenarnes@) a8 s auoercokceseeoun aud 137 

58. Breeding-basin of Myla resinifictrix: (A) side view, and (B) 
OOM) (Chien TMG) 2 oo oe Soe Ue bo od od 5544506 voueS a 158 

59. Breeding-basin of Hyla resinifictria, seen from above ..... oe ltd 

60. Diagram of the Pattern of the Blotched Tabby (Felis catus) .. 154 

61. Aorta of Tiger, showing several aneurysms ............ So te. LEG 

G2wAZyeos vems.ol Cenvicapia Donota. ses ee ass ia- Seetatiel to atniolels 186 

Gay Aayeos veins of GarellaneiGhone wniiiiess “ii. - its lreers iets sa 187 

64. Azygos veins of Phacocherus ethiopicus.... eceee snes 189 

65. Azyvos veins of Tragulus menwnnd). 0.0605. o ce tweens 191 

66. Hyrax capensis; two different arrangements of azygos system. 194 

Gi Azye0s veins Of Lrvnaceus agus) J... o-tiriansr ais 196 

GSH AZygosiOn Ge ANCrOcuba, Wn acto a twle SIP) « di-siewinle: «Weyl tae 199 

69. Azygos of Crossarchus ohscurus .......+... siiaeval avec eens .. 200 

70. Azygos veins of Halmaturus bennett .........0.0..20 ee cease 203 

71. Azygos veins of Phascolomys mitchellt ........0.ssecvevees 204 

(Pe ENA) veils Ol JESUP INGEN. 1) 5 oo BD bens au eno uo a Seo 211 

73. Lateral view of the thoracic region of newly born Myopotamus 
COU PU wa « sohansrent eee mtcR Neier dee) a Seater et Nelen ee tiedeine caeF. Meena a Sr alsa 214 

74. Hind limb at Diangimhodor Peas ee te eaeen ee eng Samia 4) DO 

“ow blind limbs on Rhamprorymenuss Wiz. aeiee atest! ths eis 225 

AGsullingdslim'a Ob Pia acany ism ose Rene ars ae ait retasts 226 


(i odind timbyot Wyetodactylus 0)... ee. 227 
(37 Hindllimbsok Hipposiderus —)......). eee net ee 30 
(OmEindlimbrot Ornichomiumus sso 0. 2) se. ee eee eye, On 
80. Hind limbs of Dipus (left) and Alactaya (right) ............ 232 
ily [altos Whirl) ore UA ACA COED Jobb bobcs cua bagdsonassdansen 233 
82. Hypothetical reconstruction of a running “ Pro-Avis” ...... 235 
85. Right mandibular ramus of Elephas antiquus (?) bearing two 
lower:molars’ Zan. ¢. aaa Mee es oct eee 245 
84, Platythelphusa armata, large male ...........00-000ceecaes 269 
80: Micracketusicollctiy a nee ee ee, ee ee 278 
BONUMicrochelusiZulicnsisi ie. eh anes eaten 280 
Oe Helis maniiean scat are eee 6 cB rar 55 300 
88. Skull of Felis manu!, view from above .............0000008 304. 
Go” Skullloterelsunanulssideicw eae ee ee 305 
SO Barbus polylends: 1a a Geay Ce amen cerns 2 ern ots RE 308 
Oy Gonbusrelephuntis® hoe). (en eee ee oe OO, 
92, Palmar surface of hand and foot of Me Halonney VS WOBHED eens o 326 
93. Part of dorsal musculature of Wegalophrys nasuta............ 302 
94, Some of the dorsal muscles of Rana gunpyt ...............- 333 
95. A dissection of Pelobates fuscus, to show large cesophageal 
sheet ot thestransyerselich see nnn e cn 304 
96. Some of the dorsal muscles of Ceratophrys orndta .......... 336 
ine klvoid tof Megalophnys nasutcmne teen 341 
98. The cesophageal sheet of transversalis muscle in Megalophrys 
OSES 65 85 ce MONT Sb B1e CRE RE RT En ete 346 
99, Liver and adjacent viscera of Rana guppyt ....... cee cee ees 349 
OOO viductotMegalopinysacasuta mee ee ee ee 350 
101. Lateral aspects of the skulls of :—a. Gymnorhina organicum. 

b. Laniarius poliocephalus. e. Sayornts cineracea ........ 355 

. Ventral aspects of the skulls of :—a. Artamus leucogaster. 

b. Sayornis cineracea. cc. Terpsiphone. 4d. Pitta baudi. 

e. Vireolanius leucotes. {. Vireo olivaced ...............- 362 
103. Ventral aspects of the skulls of :—a. Zanius collurio. b. Bas a- 
disea raygiana. c. Gymnorhina organicum................ 363 
104. Phylogenetic tree indicating the probable relationships of the 
Passeres Oligomyodii and Diacromyodii .................. 377 
105, Head of the Addo Bush, or East Cape Elephant (Hlephas 
G/PUCANUS ICAPENSIS) «2. e714. Se ee, 383 
106. Head of Female West Cape Blephant (Elephas africanus 
SOU EES EOE AS DORE EM RA. fs Acs 5 63-0. .n:6.0 080-Ao RE 385 
107. Head of Male West Cape Elephant (lephas africanus 
OUR ES) PRE AIAN Pee TR AMADA IN cc rh cin Gobioes Ad Chere 386 
108. Head of Male Mashonaland Elephant (Elephas africanus 
BELOUSL) a, SN, Fake Ie SORES ARE eee ree tere emt th 387 
109. Right Ear of Male Mashonaland Elephant (Elephas africanus 
SCLOUSD La ee ha Shia ence La ee ane: 388 
110. Head of Male Elephant from Swaziland .................. 389 
111. Right Ear of the Congo Elephant (Zlephas africanus cyclotis?). 390 


112. Head of a Male Elephant from North-west Rhodesia ........ 391 

113. Head of Male Elephant from Fort Manning, N.E. Rhodesia 
(Hlephas afiracanus knochenhawert?) .....5..g..-+-s02++-- 392 

114. Head of Male Elephant from the Aberdare Mountains, British 
Wast, Autica rants: os s)s ac naeonrame Renan rience a oth sald ~- 393 

115. Head of Male Elephant from the Lake Rudolf District (Zlephas 
AFTICONUS COVENGISHE)) io an cheEd SOTA ames eS roca, 8 ett 394 

116. Head of the “Queen’s Elephant,” an immature Male Sudan 
Hlephant, (sage ed 5) ete eR eer ae eerie. 396 

117. Right Ear of a Male Sudan Elephant (Llephas africanus 
OCU OLISI) Ys, she sie eons So Whey lag sno aah ees ETA RI La NRE RS = 397 

118. Right Ear of the North Somali Elephant (Zlephas africanus 
OUUCAIST Nin ae sales os osle sah sietanoneiearaiat eGR aa ele euSASRDs ean heen 398 

119, Head of “ Jumbo,” the male West Sudan Elephant formerly 
belongineytonbhne: Socte tyes tae ieee sie eerie 400 

120. Front view of the Skull of the Sudan Elephant (Llephas 
A ILC ANUS! OMYOLUS) aise Mea sbeu Masses 2 isn, PARENTS © eich eta oe 40] 

121. Front view of the Skull of the Albert Nyanza Elephant 
(Hlenhasyajnicaniusialbetersts) ei een eel ae eae 402 
122. Spermathecal sac of Polytoreutus ruwenzorii ......- +02 eee 418 
128. Yerminal male organs of Polytoreutus granti .......0. 2.005. 42] 
124, Spermathecal sac of Polytoreutus grant? ...............44. 422 
125, Spermathecal apparatus of Newmanniella ruwenzori.......... 426 
126. Ventral view of Eminoscoler ruwenzorat ... ........+-.-+- 498 
127. Terminal male organs of Lminoscoler ruwenzortt ...........5 430 



Monodonta (Insecta)...... 69 | Tipuliforma (Insecta) ......... .. 







Paces 1-236. 


JUNE 1907. 



ae —Price Tiuelve Shilings] ERY 

aGenal Musee 


1907, pp. 1-236. 

January 15, 1907. 
The Secretary. Report on the Additions to the Society's Menagerie during the months of 

November and December, 1906 .......- eee cece cece ee erence Sistele se ueuvenne wistnicuste 1: 

Mr. Oldfield Thomas, F.R.S., F.Z.S. Description of a new Monkey from He Ituri 

Forest. (Plate 1.) 2... cect eee cect ce cece ees cece ec cece eas essne eens stones 2 
1. On a Collection of Mammals made by Dr. Yassal in Annam. By J. Laws Bonnorg, 

M.A., F.LS., F.Z.8. (Plate II.) ..... PEPE S Arias) ah eee heen Lay AC Demo ri. & 3 
2. On the “ Bleating” or “ Drumming ” of the Snipe (Gallinago cwlestis). By P. H. Baur, 

TRAV SAHIN nia epee stalcde 10 gietei is en lace eh amie aoe Re teal emieieieh ty. Uo tate datevale 6 whe lavas feoe Bian iiee ls 
3. Contributions to the Knowledge of the Systematic Arrangement and Anatomy of certain — 

Genera and Species of Squamata. By Franz EB. Bepparp, M.A., F.R.S., Prosector to 

bE OCI by sa < vere ia allo eis cie >'» Sie vetets sie eh ee ae a eemar ea aisle Notcetarets A Hatem 14 35 
4. A List of Moths of the Family Pyralide collected by A. EH. Pratt in British New Guinea 

in 1902-3, with Descriptions of new Species. By Gnorczk H. Kenrick, ¥.Z8. 

(Pints WUT, DV) Fie vies otaemie esanclse ale een retrere Pstarete AE Fy Ae a ES oh 68 
5. On some new and insufliciently known Species of Marmoset Monkeys from the 

Amazonian Region. By Prof, Dr. Bui A. Gorrnr, C.M.ZS., Director of the Para 

Museum 2.0.6 cece eee te ee cece reece eet ee tnt et ene cnet eee cette es Arata 88 

February 5, 1907. ; 

Myr. F. Martin Duncan. Cinematograph exhibition of Animals in the Society’s Gardens 

and other Zodloridal Subjects. ceo POH es ihe wo aera 2 wiahinten sian ate acsiate siete tate tetele 100. 
Mr. Oldfield Thomas, F.R.S., F.Z.S. Exhibition of a collection of Mammals and Birds 

from the Islands of Saghalien and Hokkaido ........6..cseeeeeee SR sna c lee SOU 

Contents continued on page 3 of Wrapper. 



Tuts Society was founded in 1826 by Sir Sramrorp Rarrtes, 
Mr. J. Sasrnz, Mr. N. A. Vigors, and other eminent Naturalists, 
for the advancement of Zoology and Animal Physiology, and for the 
introduction of new and curious subjects of the Animal Kingdom, 

and was incorporated by Royal Charter in 1829. 



George A. Bovnenenr, Ese., | P. Caarmers Mrrcnert, Ksa., 

F.R.S., Vice-President. | EAMG iD USton, IDRIbRIB 5. aderatSin. 
Joun Rosz Braprorp, Ese.,M.D., | Secretary, 

D.Sc., F.R.S., Vice-President. || W. R. Ocinyie-Grant, Ese. 
F. Dawrrey Drewirr, Ese, || Aubert Pam, Esa. 

vAG,, M1). | s 

M.A., M.D Ki. Lorr Parties, Ese. 

CHartes Drummonp, KEsa., 

P | Str Patrick Prayrarr, C.LE. 
Treasurer. | 
Srr Epwarp Durano, Br., C.B. Tun Hon. Watrer Roruscatzp, 

F. Du Cane Gopman, Hse, | D.Se., M.P. 
D.C.L., F.R.S., Vice-President. || Howarp Saunpers, Hsq., Vice- 

Josep Jackson Lasrer, Ese., | 
M.A., F.RB.S. | Davin Sera-Surra, se. 

Srr Epmunp Grows Lopur, Br., | OLDFIELD Tomas, Hse, F.R.S. 
Vice-Presedent. | A. Truvor-Barryg, Kse., M.A. 

Pror. Epwarp ALFRED Mincury, || Henry Woopwarp, Hse., LL.D. 
M.A. F.R.S., Vice-President. 


The Society consists of Fellows, and Honorary, Foreign, and 
Corresponding Members, elected according to the By-Laws. It 
carries out the objects of its foundation by means of the collection 
of living animals at Regent’s Park, by its Library at 3 Hanover 
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The Office of the Society (3, Hanover Square), where all 
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open from Ten till Five, except on Saturdays, when it closes at 
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The Library, under the superintendence of Mr. F. H. Waterhouse, 
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The Meetings of the Society for General Business are held at the 
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The Meetings for Scientific Business are held at the Office twice 
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The Gardens in the Regent’s Park are open daily from Nine o’clock 
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The Prosectorium for Anatomical and Pathological work at the 
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Prosector, assisted by Dr. ©. G. Seligmann, Pathologist to the 


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3 Hanover Square, London, W., 

June, 1907. 



TuEspay, JANUARY 15 Turspay, May .... 7 and 28 
¥y Feprvary 5 and 19 if JUNE ees 
és March .. 5 «,- 19 56 November 12 and 26 
- AUER oy SN oy 28} ., DecremBer 10 

The Chair will be taken at half-past Eight o'clock in the Evening 




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The following is a complete list of the publications of the 
Society already issued. 

[ June, 1907. ] 


4to. 16 vols. and Index. Enice te Price to the 
ellows. Public. 
Vol. _I., containing 59 Plates.... (1833-85) .... £3 13 6.... £4 18 Of 
Ree he 7... (188A) ae MO ONS 6 Gt 
Gaplliee a VARNES Thee ©. (1GAD EMO) pe ancien, 
7 Tai ii gale (isb1262) 6 2 0 Ss 2 cor 
MEN VG j-!.., . KIRC266)) 2: ts ae ee TOPO 
Pavel, *5 tn Gea e CKO oad Lk SB © , 15 0 O 
2 Vl ASE AO... 1319 © 
peal T es | iy go 1). (ere ot AO ie bs) ee SO 
EI, meme (Ge) os eT Gays 1G 9 0 
G Jone SCA) sooo: O 3, le 7 O 
Index, Vols, ATS aXat td vs viele ante EES (O)eotn Of Coos5 OO 
Vol. XI., containing 97 SBIR: 3 (U880=85) a OD Oa al eG uO 
i X11, yy | OD 2 yy? oo (esGH80) sogn O 8 O; i 4 
xa, 5 Gs 4, (RSIS) 6.8 18 4) allen 
axe 2 AT Ne a UB OCLOS) germs 45 THO 7 0 0 
Lexa P52 9 2 eosligon) (2175 156 eran 
ON ee CLIC eee, 7 AO 
GV see wl: as, ae on (Anes ISOS) 55 i a Gy 110 0 
UX oy 2 one ¢ Boh, vor Aup OOS) gate 0 13 36 | 12 wonemm 
CH ne ae ea (ORC ONS ON a LO o 
Seyi, ae, 88) of (Oct MOOD) ee el 72 46... gel NO 
OVI Ne Ree ERG act (Decwl905) = x0 115 0 | somo 
oy le ote OF by IE oa (CGtin ITS)o 0.5 Ib BG 110 0 


LONDON. 8vo. 2 vols. (Letterpress only). rice te Eley eone 
Raney Ilo ISO. vO, ByOs “soasooscasonce 4s. 6d. 6s. 
DRS 2 ety aye Meco asie rte cenconeto ors As, 6d. 6s. 

9 i) 

8vo. 15 vols. (Letterpress only) and Index. (First Series.) 

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(January to April, 1907.) 

January 15, 1907. 

Dr. J. Rosz Braprorp, F.R.S., Vice-President, 
in the Chair. 

The Secretary vead the following report on the additions that 
had been made to the Society’s Menagerie in November and 
December 1906 :— 

The registered additions to the Society’s Menagerie during the 
month of November were 173 in number. Of these 105 were 
acquired by presentation and 23 by purchase, 35 were received on 
deposit, 3 in exchange, and 7 were born in the Gardens. The 
total number of departures during the same period, by death and 
removals, was 177. 

Amongst the additions special attention may be directed to :— 

An adult male Mandrill (Papio maimon), the first full-sized 
example of this species exhibited in the Gardens, deposited on 
Noy. 30th. 

A young female Hippopotamus (Hippopotamus amphibius) trom 
the Niger, purchased on Novy. Ist. 

A Persian Stag (Cervus maral), presented on Nov. 13th by 
Mr. Carl Hagenbeck. 

A Kashmir Stag (Cervus cashnuriensis), presented on Noy. 22nd 
by H.G. the Duke of Bedford, K.G., P.Z.8. 

A Collection of 47 Birds, including, among other interesting 
species, a Toucan (Awlacorhamphus sulcatus), new to the Collection, 

Proc. Zoou. Soc.—1907, No. I. 1 


anda Sun-Bittern (Hurypyga helias) from Venezuela, presented 
on Nov. 27th by Capt. A. Pam, F.Z.8. 

The registered additions to the Society’s Menagerie during the 
month of December were 150 in number. Of these 67 were 
acquired by presentation and 16 by purchase, 61 were received on 
deposit, 2 in exchange, and 4 were born in the Gardens. The 
total number of departures during the same period, by death and 
removals, was 207. 

Amongst the additions special attention may be directed to :— 

A pair of Siberian Dholes (Cuon alpinus) from Thian Shan, 
received in exchange on Dec. 2nd, new to the Collection. 

A Cape Hunting-Dog (Lycaon pictus) from South Africa, pur- 
chased on Dee. Ist. 

An Addax Antelope (Addax naso-maculatus) from North Africa, 
presented by H.G. the Duke of Bedford, K.G., P.Z.S., on Dec. 18th. 

A Bubaline Hartebeest (Alcelaphus bubalinus), and a hybrid 
between Pére David’s Deer (Hlaphurus davidianus) and the Red 
Deer (Cervus elaphus), deposited on Dee. 29th. 

A new Monkey from the Ituri Forest. 

Mr. Oldfield Thomas, F.R.S., F.Z.S., exhibited a Monkey which 
had been obtained in the Ituri Forest, Upper Congo, during the 
recent Ruwenzori Expedition, and gave the following description 
of it :— 


Abstr. P. Z.S. 1907, p. 1 (Jan. 22, 1907). 

A member of the campbelli-mona group, but not darkened on 
the posterior back and hind limbs, and with a very sharply con- 
trasted white belly. 

Upper surface of head and neck olive-grey, the usual light frontal 
baud present but not conspicuous. Back dark grizzled chestnut- 
brown (nearest to “burnt-umber” of Ridgway); colour of rump 
not darker, but, on the contrary, passing gradually into the 
paler tone of the hips and hind legs. Under surface from chin 
to anus, and inner sides of limbs to wrists and ankles, clear 
creamy-white, very sharply defined from the darker colour, not 
only on the limbs, as in campbelli and others, but also along the 
flanks, where the white rises nearly halfway up the lateral aspect 
of the animal. Ears with short yellowish tufts rising from their 
inner surfaces. Outer sides of fore limbs deep black from 
elbows. Hind limbs grizzled yellowish olive, lighter than the 
back, down to and including the ankles, the metatarsals and toes 
black. Tail indistinctly blackish above at base, then dull greyish 
white for two-thirds its length, darkening again to black on its 
terminal third. 

* [The complete account of the new species described in this communication 
appears here ; but since the name and preliminary diagnosis were published in the 
‘Abstract,’ the species is distinguished by the name being underlined.—Eprror. | 

SZ SIO yvealk 1. 



Bale & Danielsson a imp. 

J. Smit hth. 


= i 

de 239 USO, JENA, 

H. Goodchild, del et ith. 

Huth, amp ; 


Dimensions of the type, measured in the flesh :— 

Head and body 501 mm.; tail 850; hind foot 155; ear 40. 

Skull— greatest length 105 mm., basal length 75; breadth of 
brain-case 55; length of upper cheek-tooth series 23. 

Hab. Ituri River, between Mawambi and Avakubi, Upper 
Congo ; alt. 8000’. 

Type. Adult male. B.M. no, Original number 184, 
Collected 23 October, 1906, by R. HE. Dent. 

This handsome Monkey 1s most nearly allied to the W. African 
C. campbelli, but differs by its grizzled olive-yellowish instead of 
black hind limbs, the absence of black on its posterior back, its 
more or less greyish-white tail, and by the high and sharply 
defined line separating the colours of the flanks and belly . 

The following papers were read :— 

1. On a Collection of Mammals made by Dr. Vassal in 

Annam. By J. Lewis Boynore, M.A., F.L.S., F.Z.8.* 
[Received November 16, 1906. ] 
(Plate IL.7 and Text-figures 1, 2.) 

The British Museum has recently acquired a most interesting 
set of Mammals from Annam, collected by Dr. Vassal. The 
collection contains examples of some twenty-five species, of which 
five are new to science, while several of the others add consider- 
ably to our knowledge (still very limited) of the fauna of the 

Since the collections made by MM. Pierre and Mouhot over 
half a century ago, practically no fresh material has reached 
Europe from that locality. As would therefore be expected, 
many of the forms are undescribed, and there is little doubt that 
with further material many of the forms at present included 
under existing names will prove to be subspecifically distinct. 

The collection is, perhaps, too small for any generalisation on the 
fauna of Annam, but its affinities seem if anything to tend towards 
China rather than the Matay Peninsula, and it is especially note- 
worthy that it differs considerably from the fauna of Siam. Lest 
I am misunderstood, I may as well point out that by “fauna” I 
am not referring to the presence or absence of certain genera, 
but rather to the fact that the local forms of widely spread 
species approximate rather to the Chinese than to the Malayan. 
To give some examples:—The Porcupine is Anderson’s Hystria 
yunnanensis, not H, groter from the Peninsula. The Petaurista 
is Anderson’s P. yunnanensis, and not P. lylei, mihi, from Siam. 
The new Zupaia described has its affinities with 7. chinensis and 

* [The complete account of the new species described in this communication 
appears here; but since the names and preliminary diagnoses were published in the 

* Abstract,’ such species are distinguished by the name being underlined.—Ep1rTor. | 
+ For explanation of the Plate, see p. 11. 

4 MR. J. L. BONHOTE ON [Jan. 15, 

not with 7’. ferruginea. On the other hand, the Paradoxurus is 
apparently identical with a form described by me from the 
Peninsula. Another point of interest as showing a probable 
double origin for this fauna, is in the occurrence at the same place 
of two subspecies of Sciwrus macelellandi—one, S. m. rodolphi 
A. M.-E., showing very obvious affinities with S. m. barbei of 
the Peninsula; the other, S. m. maritiémus mihi, which is in- 
distinguishable from the type, which came from China. It must, 
however, be remembered that this last is only represented by a 
single skin, and it might possibly have been brought down on a 
ship and escaped. 

Lastly, attention may be called to a new species of Vycticebus, 
which is in many respects intermediate in its characters between 
Nycticebus and Loris. 

As regards synonymy, I have followed my usuai custom, 
namely, to give the original reference and a few of the other more 
important ones, which, if referred to, will be found to contain a 
practically full synonymy. 


Semnopithecus nigripes A. M.-Edw. Nouv. Arch. du Mus. 
WO Talo JUNIE joe Wy iol WCherAbys lelhyilng di, As So 185 Sschiny, Ess IO: 
p- 11 (1875); Anders. Zool. Res. p. 41 (1879). 

a. 6 ad. Bali, alt. 250 m., 10th Nov., 1905. 

This is an extremely fine example of this scarce species, 
agreeing very well with the published descriptions. 

a,b. 2. Nha-trang, 30th Oct., 1905. 

Two very young specimens of a species of Presbytes, unfor- 
tunately too young for identification. 

Nycricesus pyemaus Bonh. (Plate II.) 

Abstr. P. Z.S. 1907, p. 2 (Jan. 22, 1907). 

Very small, about half the size of . cowcang* Bodd. The 
hair is wavy on the body and of a very fine silky texture. 
General colour of a uniform orange-rufous, showing no sign of 
any dark line down the back or on the head. The under parts, 
hands, and feet are lighter in colour and have a silvery-grey 
appearance. There is a bare space round the eyes, the muzzle 
and lips are white, and a white stripe runs up from the nose 
between the eyes to end abruptly on the forehead. The ears are 
of moderate size, uniformly rounded, and very sparsely covered 
with hairs. The tail is a mere stump. 

The skull in its general outline agrees fairly well with that of 
NV. c. cinereus from Cochin China, although it is, of course, very 
much smaller. In its main characters also it shows no very 

* Messrs. Stone and Rehn have pointed out (Proc. Acad. Nat. Sci. Phil. 1902, 
p. 138) that the name tardigradus belongs to the Slender Loris “ L. gracilis,” and 

that therefore Boddaert’s name must stand for the Slow Loris usually known as 
N. tardigradus, , 


distinctive points. The molars, however, are conspicuously 
different, and enable this species to be easily diagnosed. In 
AV. cinereus the first molar is the largest, and the last or third 
molar is small and almost quadrilateral in shape. In the present 
species, however, the second molar is the largest, while the third 
molar is triangular in outline and not very much smaller than the 
first molar. In the lower jaw similar conditions obtain, the three 
molars are all subequal, the third being slightly the largest, 
whereas in iV. cinereus the last molar in the lower jaw is very 
markedly smaller than either of the other two. 

Dimensions of type from skin (approx.). Head and body 
190 mm.; tail 10. 

Text-fig. 1. Text-fig. 2. 

Text-fig. 1.—Palatal view of skull of Nycticebus pygmeus. 
Text-fig. 2.—Lateral view of skull of Nycticebus pygmeus. 

Skull. Greatest length 46 mm.; basal length 38; palatal length 
17-5; zygomatic breadth 27; interorbital breadth 3; greatest 
breadth of brain-case 25; length from palate to lower margin of 
foramen magnum 17:5; breadth of basioccipital at its anterior 
end 3°7; length of molar series 14. 

Hab. Nha-trang, Annam. 

Type. B.M. 3. Collected by’ Dr. Vassal on the 
13th Nov., 1905. 

The small size and peculiar character of the teeth will prevent 
this species from being confounded with any others at present 
known to exist. Only a single specimen (the type) has been sent, 
which is quite adult but not old. It may be noted that in some 
respects the teeth tend to approach those in Loris, in which the 
second upper molar is larger than the first. In the shape of the 
premaxilla also the present species shows a tendency, albeit very 
slight, to approach Loris by showing its flat surface laterally 
instead of anteriorly. Externally the blaze between the eyes and 
its small size are features belonging to Loris, but in the length of 
its limbs and general build it is a true Vycticebus. 


MR. J. L, BONHOTE ON [Jan. 15, 

FELIS sp. ? 

a. Flat skin with no data of a Cat belonging apparently to the 
Felis bengalensis group. 


Viverra megaspila Blyth, J. A. 8. B. xxx. p. 331 (1862). 

a. Nambon, Annam. 

A fine adult specimen with well-marked and clear-cut spots. 


Viverra malaccensis Gmel., Linn. Syst. Nat. i. p. 92 (1788); 
Gray, P.Z.S8. 1861, p. 136. 

Viverricula malaccensis (Gmel.), Bonhote, Ann. & Mag. N. H. 
ser. 7, vol. 1. p. 118 (1898). 

a. Imm. No data. 


Paradoxurus minor Bonh. Fasci. Mal., Zool. i. p. 9 (19038). 

a,b. 9 imm. Bali, Annam, 250 m., 10th Nov., 1905. 

These are both very young specimens, which agree closely with 
the type. 


Herpestes exilis Gerv. Zool. de la Bonite, p. 32 (1841); Gray, 
P.Z.8. 1864, p. 555. 

Herpestes javanicus (Geoft.), Anders. Zool. Res. p. 185 (1879). 

Herpestes rutilus Gray, P.Z.8. 1861, p. 136. 

Calogale rutilus Gray, P.Z.S. 1864, p. 561. 

a,b. 9. Nha-trang, 26th Dec., 1905. 

I have carefully compared these specimens with some from 
Siam and others from Cochin China, among them Gray’s type 
of H. rutilus. The Cochin China and Annam specimens are all 
very like each other, and differ in their much redder colour from 
Siamese specimens. They also differ in their much deeper colour 
from Javan specimens. Gervais’s type of H. ewilis came from 
Cochin China, and as his description agrees fairly well with these 
fresh specimens, his name of H. exilis, which antedates Gray’s, 
must stand. 

The skulls of H. emilis, although very similar to those from 
Siam and Java, are larger and more robust. The Siamese animal 
is apparently intermediate between H. birmanicus and H. ewilis. 

The following is a description of the present specimens :— 
General colour rufous, punctulated with white. Each hai is 
black, with three or four buff or rufous annulations. The distal 
annulations and generally the tip of each hair are rufous, while 
along the centre of the back, the head, cheeks, and tail these 
rufous annulations are deeper in colour and more marked, causing 
the animal to appear quite red along those areas. The under 
parts are more sparsely covered with hairs and the annulations 

yellowish rather than rufous, except under the chin and at the 
root of the tail. The hairs of the tail, more especially under- 
neath and at the sides, have long rufous tips. 

Dimensions (of Nha-trang specimen, ad. 2 in flesh). Head and 
body 364 mm.; tail 284; ear 28. 

Skull. Greatest length 78 mm.; basal length 75; zygomatic 
breadth 39; palatal length 41; greatest diameter of carnassial 8. 


Helictis pierrei Bonhote, Ann. & Mag. Nat. Hist. ser. 7, vol. xii. 
p. 592 (1903). 

a. Imm. Nha-trang, Annam. 

6b. Imm. skull only. 

The single skin and skull are too immature to show the dis- 
tinctive characters to any marked extent. 

TuPAIA Concotor Bonh. 
Abstr. P. Z. 8. 1907, p. 2 (Jan. 22, 1907). 

Similar in general colouring to Zupaia belangeri. The whole 
of the upper parts are of a uniform grizzled greyish-green, each 
hair being dark at its base and having one or more buff annulations 
and a dark tip. One of the most distinctive features is the absence 
of the neck-stripe, so universal among the other species of this 
genus. An extremely faint trace of it only is to be made out on 
the shoulders, but so faint is it that unless special search be 
made it is liable to be overlooked. The tail, which is markedly 
distichous, is concolorous with the upper parts, and extremely 
thick and bushy. The under parts are somewhat sparsely clothed 
with hair; the chin, throat, and breast are uniformly yellow, 
while on the other portions the hairs are annulated as on the 
upper parts. The bases of the hairs on the under side of the tail 
are light. 

Skull. In its general character resembles that of 7’. belangeri; it 
is, however, slightly larger and witha longer and narrower snout, 
in other respects it does not show any marked features. 

Dimensions of type (from skin). Head and body 220 mm. ; 
tail 140; ear 15; hind foot 45. 

Skull. Greatest length 54 mm.; basal length 47; zygomatic 
breadth 29; palatal length 27; breadth of skull immediately 
behind postorbital processes 17. 

Type. B.M. o ad. Collected by Dr. Vassal, 22nd 
March, 1906. 

Hab. Annam. 

Although very closely allied to Tupaia belangeri this species 
may easily be distinguished by its larger size, much thicker tail, 
and the absence of the light neck-stripe. Two specimens agreeing 
in allrespects were brought back by Dr. Vassal. Tupaia chinensis, 
described by Dr. Anderson from Yunnan and which is found in 
Siam, is rather smaller than 7’. belangeri and consequently quite 
distinct from the present form. 

8 MR. J. L. BONHOTE ON [Jan. 15, 


Tupaia frenata Gray, Ann. & Mag. Nat. Hist. ser. 3, vol. vi. 
p. 217 (1860). 

Dendrogalefrenata Anders. Zool. Res. p.110, pl. 7. figs. 20, 21(1879). 

a, 0. 

Two very typical examples. 


Vespertilio sphina Vahl, Skrivter af Naturhistorie-Selskabet, 
Ate Band, lste Heft, p. 123 (1797). 

Cynopterus sphinw (Vahi), Bonh. P. Z. 8. 1902, vol. i. p. 38; 
id. Fasci. Mal., Zool. pt. i. p. 14 (1903). 

a. 6. Nha-trang, Annam, 13th Nov., 1905. 

ScoroPpHiLus KUHLII Leach. 

Scotophilus kuhli Leach, Trans. Linn. Soc. xiii. p. 71 (1822). 

a,b. $. Nha-trang, 10th Oct., 1905. 


Pteromys yunnanensis Anders. Zool. Res. p. 282, pl. xxi. (1879). 

a. 6 ad. Bali, Annam, 150 m., 10th Nov., 1905. 

This individual agrees very well with Dr. Anderson’s description 
and plate (quoted above), and I have no alternative but to place 
it under his name. At the same time it should be noted that the 
typical locality of P. ywananensis is considerably to the north and 
that another form of this same species, ?. lylei mihi, is found 
in Siam. Jt would therefore appear as if the present race was 
in reality a Chinese form and that Annam and Yunnan form its 
western limit. Except for the parachute the hairs of the whole 
of the back in this individual are tipped with white, but not 
sufficiently so as to conceal the chestnut ground-colour. 

P. lylei is much darker in general coloration than this species 
and the anterior portion of the outer side of the ear is a bright 
and pure chestnut. 

P. yunnanensis and P. lylei belong to a large group, of which 
there are many geographical forms. Until, however, the group is 
worked out as a whole, it is best to retain them under binomial 
names, but it should be borne in mind that they are merely 
geographical forms of a large and widely distributed species. 


Sciurus griseemanus A. M.-Edw. Rev. Zool. 1867, p. 195; 
Bonh, Ann. & Mag. Nat. Hist. ser. 7, vol. vil. p. 274 (1901). 

a-d.3 3, 9. Nha-trang, Annam, 26th Dec., 1905. 

e. Nha-trang, Annam, Nov. 1905. 

f-k. 33,2¢. Hoah Khat, Annam, 26th Dec., 1905. 

In. 26, 9. Ninh Hoa, 25th Dec., 1905. 

o-p. 6. Bali, Annam, 250 m. alt., 26th Dec., 1905. 

The present species and S’. lewcopus* of Gray have hitherto been 
confounded and considered as one and the same species. The 

* Macrowus leucopus Gray, Ann. & Mag. N. H. ser. 3, xx. p. 282 (1867). 


present series, however, shows that they are really quite good and 
distinct species. The most obvious difference is in the colour of 
the under parts. In S. griseémanus they are deep chestnut and the 
line of demarcation between the upper and under parts is sharply 
divided. In S. leweopus, on the other hand, the colour of the 
under parts is of a pale rufous buff, which shades gradually into 
the grizzled grey of the back. M. Milne-Edwards in his original 
description of S. griseimanus distinctly states that the colour 
of the under parts is deep chestnut, though females and young 
males are sometimes considerably lighter. This enables us to 
fix M. Milne-Edwards’s name on the chestnut-bellied form with- 
out hesitation. In the present series the colour of the under 
parts is very deep chestnut and shows but little variation ; the 
two examples from Bali, at an altitude of 250 metres, are, how- 
ever, much lighter below, and it may be that these lighter 
individuals represent a mountain race of S. griseimanus, but our 
material is at present too scanty to settle that question. 

S. leucopus differs still further from S. grisetmanus in the 
annulations on the hairs of the back being yellower and not of 
such a clear grey, thus giving the animal a darker appearance. 
The colour of the under parts also extends over the outer sides of 
the limbs and is especially noticeable on the thighs. 


Abstr. P.Z.S. 1907, p. 2 (Jan. 22, 1907). 

General colour above similar to that of S. lewcopus, but darker 
(see preceding species). Hach hair is very dark, with three or four 
yellowish annulations. The tail, which is indistinctly barred, is 
similar in colour to the body, but “4he amnulations are of a slightly 
deeper tint. Hands and feet dirty yellowish white grizzled with 
darker. Under parts and inner sides of the limbs “pale reddish 
buff, with the exception of the chin and throat which are grizzled 
like the back. 

The skulls available are so imperfect that a description 1s not 

Dimensions of type (from skin). Head and body 190 mum. ; 
tail 175; ear 17; hind foot 52. 

Hab. Ninh Hoa, Annam. 

Type. B.M. Collected on the 10th Nov., 1905, by 
Dr. Vassal. 

This species is easily distinguished from S. lewcopus typicus by 
its darker colour above, grizzled hands and feet, and by the outer 
sides of the limbs being similar in colour to the rest of the upper 
parts. Whereas the typical .S. lewcopus is greyer, the outer sides 
of the limbs are buff, and the hands and feet pure yellowish white. 


Sciurus macelellandi maritimus Bonh. Ann. & Mag. Nat. Hist. 
ser. 7, vol. iv. p. 51 (1899). 
* Since the reading of this paper it has been pointed out to me that the name 

“< fumigatus ” 1s preoccupied, haying been used by Gray in 1867. I therefore propose 
to rename this squirrel Sciwrus vassali. 

10 MR. J. L. BONHOTE ON [Jan. 15, 

a. &. One specimen, 10th Nov., 1905. 

A single example of this Chinese race has been brought back ; 
it resembles the type closely and in all respects. The occurrence 
of this form in Annam is certainly surprising; it may, however, 

pee aoe , 
prove to range along the whole S. Chinese coast, while S. rodolphi 
inhabits the higher ground; on the other hand it might have been 
brought over on a ship and escaped. 


Seiurus rodolphi A. Milne-Kdwards, Rev. et Mag. de Zool. xix. 
p. 227 (1867); id. Rech. Mamm. p. 162 (1871). 

Sciurus macclellandi rodolphi A. M.-E., Bonh. Ann. & Mag. 
Nat. Hist. ser. 7, vol. iv. p. 54 (1899). 

a-c. Three specimens. 

These are very typical specimens. The light stripes are of the 
same width throughout, and have a tendency to a deeper and more 
rufous tinge on their anterior portion. The median dark stripe 
tends to become divided down the centre by a brownish grizzled 
stripe; the length and extent of this latter stripe seem to be very 


Sciurus berdmorei Blyth, J. A.S. B. xvii. p. 63 (1849); Anders. 
Zool. Res. p. 261 (1879). 

Funambulus berdmoret (Blyth), Bonh, P. Z. 8. 1901, vol. i. p. 56. 

a, b. Bali, Annam, 10th Nov., 1905. 

S. mouhoti Gray was merely described on a seasonal form of 
this species, as I have already pointed out. 


Abstr. P. Z. 8. 1907, p. 2 (Jan. 22, 1907). 

Similar to /. rufigenis typicus, but the general tone of colour 
very much deeper. Colour above very dark brown, finely grizzled 
with buff. The whole of the thighs suffused with deep chestnut, 
which is not the case in the typical form. Sides of face chestnut, 
rather deeper in tint than in the typical form, and the same may 
be said of the chestnut on the under side of the tail. Remainder 
of under parts creamy white. 

The skull appears to be of a rather stouter build, but without a 
good series it would be unwise to lay stress on this fact. 

Dimensions of type (from skin). Head and body 190 mm.; 
tail (broken) 150; ear 12°5; hind foot 43. 

Hab. Bali, Annam, 250 m. alt. 

Type. B.M. Collected by Dr. Vassal on the 10th 
Nov., 1905. 

The much darker general colour and the rufous tinge on the 
outer sides of the thighs form good distinctive characters by which 
this species may be easily distinguished from the typical race. 

Ruizomys pRuiNosus Blyth. 
Rhizomys pruinesus Blyth, J. A. 8. B. xx. p. 519 (1851); id. 


Cat. Mamim. As. Soc. Bengal, p. 122 (1863); Anders. Zool. Res. 
p. 325 (1879). 

a. Plateau of the Lung Brau, 1300 m., 30th Oct., 1905. 

This specimen is only a flat skin without a skull; Dr. Vassal 
notes that it is “‘not rare” in Annam. 

Hysrrix yYUNNANENSIS Anders. 

Hystrix yunnanensis Anders. Zool. Res. p. 332 (1879). 

a. 2 imm. Ninh Hoa, 25th Dec., 1905. 

This is a very young specimen, but has a well- developed nuchal 
crest. The skull in its general proportions agrees with Dr. 
Anderson’s description. The external characters, however, agree 
well with Swinhoe’s /7. swheristata, and, in fact, the only difference 
between these two species is to be found in the skulls. In his 
original description Swinhoe states that the skull of . suberistata 
is indistinguishable from that of MH. hodgsoni Gray, which, of 
course, is quite distinct from Anderson’s species. Unfortunately 
there are no specimens from China in the British Museum which 
would enable us to determine definitely whether there be two 
Crested Porcupines in China, or whether Anderson’s and Swinhoe’s 
species are in reality one and the same. 

Lepus vAssati Thos. 
Lepus vassali Thos. Ann. & Mag. Nat. Hist. ser. 7, vol. xvi. 
p. 425 (1906). 
9. Nha-trang, Annam, 25th Dec., 1905. 
Mr. Thomas having recently described this specimen, I need 
only refer those interested to the paper quoted above. 

Tragulus affinis Gray, P.Z.S. 1861, p. 138. 
Tragulus kanchil pierret Bouh. Ann. & Mag. Nat. Hist. ser. 7, 
vol. xi. p. 293 (1903, 1st March). 
a-c. 3. Nha-trang, 22nd March, 1906. 
aie Specimens agree in all respects with my description of 
. k. pierret quoted “above. I have used Gray’s name for this 
Hess in preference to my own, as Mr. Miller has pointed out to 
me that Gray’s 7’. affinis was chiefly based on specimens from 
Cochin China; and Mr. Miller having, previously to my paper, 
described the Peninsula form under the name 7’. ravus, Gray’s 
T. affinis became ipso facto restricted to the race from Cochin 
China: with this finding I quite agree. 


Manis javanica Desm. Mamm. p. 377 (1820); Anders. Zool. 
Res. p. 352 (1879). 

a. Dang-trang, near Nha-trang, 25th Dec., 1905. 

Nycticebus pygmeus, p. 4. 

12 MR. P. H. BAHR ON THE (Jan. 15, 

2. On the * Bleating” or “ Drumming” of the Snipe 
8 g P 
(Gallinago celestis). By P. H. Baur, B.A., F.Z.8. 
[Received November 20, 1906. | 
(Text-figures 3-9.) 

This subject has been much discussed, but the interest taken in it 
seems rather to have waned during the latter years. I think there- 
fore it would be profitable to inquire once more into this strange 
phenomenon, especially as many points require elucidation, in the 
explanation of which authorities differ considerably. I believe it 
is well known to all of us that during the breeding-season our 
Common Snipe performs certain aerial evolutions, producing at the 
same time a mysterious sound, called in various parts of the country 
bleating, humming, drumming, or whirring, ‘“‘ Meckern” in Ger- 
many, while in parts of Scotland the popular name of the bird is 
‘* Heather Bleater,” and in France “ Chévre volant.” The process 
is Shortly as follows :—The bird is seen to fly straight up to a 
height of from 60-100 feet, then, turning, to spread its tail, close 
its wings, and drop to within 20-30 feet of the ground, producing 
at the same time this mysterious sound, which has puzzled observers 
so. As to the cause of it many theories have been put forward 
by scientific ornithologists, sportsmen, and foresters. We may 
group the evidence for these theories under four heads :— 

I. The sound is produced by the vocal organs. 
II. The sound is produced by the rectrices of the tail, which I 
hope to be able to prove is correct. 
Il. The sound is produced by the action of the primaries of the 
IV. The sound is produced by the combined action of the wings 
and tail (maintained by a few observers). 

I. The first evidence in favour of this theory which I can find 
is an article by Débel (‘Jiiger Practica,’ 1783, pt. i. p. 73), who 
says that the Snipe produces at night-time, while sitting on the 
ground in a marsh or close to water, a noise which the ignorant 
would mistake for the bleating of a young goat. 

Bechstein, in 1789 (Naturgesch. Deutsch. 2nd ed. vol. iv. p. 190), 
maintains “that the Snipe makes its plaintive ery, like a goat 
bleating, with its beak and not, as has been lately affirmed, with 
its wings.” He adduces evidence of birds bleating whilst perched 
on the tops of trees. 

Hintz (‘ Naumannia,’ 1854, p. 290), from observations made 
from 1816-19, believes the sound is made by the bill. He heard 
Snipe “bleating” whilst sitting on top of withered oak-trees, first 
uttering their call-note “ pecka pecka,” and then bleating. 

Zoppritz (Ornith. Centralblatt, Nov. 1880) seems to be very 
certain of the accuracy of his observation, for he writes :—‘ Two 
years ago I published an article on this subject in a sporting 
journal, wherein I offered to pay a fine of 500 marks to the treasury 
of the Allgemeine Deutsche Jagd-schutz-verein if three umpires 
appointed by the Verein publicly declared that they were convinced 

1907. ] ‘‘BLEATING” OF THE SNIPE. 13 

that Snipe produced their bleating notes, not through the vocal 
organs, but by means of their wings, with or without help of their 
tail-feathers.” Nobody seems to have accepted the offer, and we 
understand Herr Zéppritz kept his 500 marks. Again, we find 
in the same paper :—‘ Anyone with a knowledge of mechanics or 
physics, if he sufficiently examine a dead Snipe, must be convinced 
that so small a bird, with wing- and tail-feathers so comparatively 
weak, cannot possibly produce sounds with them, which are at 
such a distance so sharply accentuated. Hence [indulge the hope 
that the adherents of the wing and tail theories will now with- 
draw their opinions and acknowledge that to err is human.” 

Lastly, in Seebohm’s ‘ British Birds,’ vol. ii. p. 244 (1885), we 
find the following:—‘‘T have listened to the drumming of the 
Snipe scores of times with the express purpose of discovering the 
mode in which the sound is produced, but must confess myself 
completely puzzled. Arguing from analogy, I should say it was 
produced by the vocal organs, and is analogous to the trill of the 
Stints and other Sandpipers. The fact that it appears to begin 
the instant the bird begins to descend induces me to think that, 
after allowance is made for the time it takes for sound to travel, 
it must really begin before the descent, whilst the bird is not 
moving very rapidly.” 

Of such is the evidence with which we have to deal. At 
the present day I trust this theory has but few adherents. Pralle 
seems to have disposed of it entirely by a note recorded in ‘ Nau- 
mannia’ (1852, pt. 1. p. 25), that on 24th March, 1846, he heard the 
Snipe utter its note, “ gick-jack, gick-jack,” while bleating. 

If. That such a sound could be produced by such feeble instru- 
ments as the rectrices seems to have been foreshadowed by 
Naumann, curiously enough by a misprint of the word “tail” for 
‘“‘ wing-feathers ” (‘ Federwild-jagd,’ von Louis Liegler, Hannover, 
1846, p. 174) :—“ It is not hard with sharp eyes (still more with 
field-glasses) to observe the quivering motion of the tips of the 
tail-feathers [the italics are my own] during each downward and 
upward flight through the air, sufficient to convince one that the 
sound is thus produced, and not from the throat of the bird. ‘The 
sound, or at least a similar one, can be produced if one take the 
primaries of certain (but not too small) birds and fasten them to 
the end of a long cane, and strikes with this, as with a sword, ina 
draught of air.” 

To which Jickel (‘ Naumannia,’ 1855, pp. 112, 113) replies and 
casts doubt on Naumann’s theory (or, rather, mistake) of the sound 
being produced by tail-feathers. 

In 1858 Mr. Wolley communicated a paper by Mr. Meves, of 
Stockholm (Proc. Zool. Soc. Lond., April 1858, p. 199), wherein 
Mr. Meves, in consequence of the misprint already quoted, was 
led in 1856 to make experiments with the rectrices of G. celestis. 

He remarked with surprise “that the humming sound could 
never be produced whilst the bird was flying upwards, at which time 
the tail is closed; but only when it was casting itself downwards 
in a slanting direction, with the tail strongly spread out.” 

14 MR. P. H. BAHR ON THE (Jan. 15, 

He then examined the tail-feathers of our common species 
more closely and found “the jist (outer feather) especially very 
peculiarly constructed ; the shaft uncommonly stiff, sabre-shaped. 
The rays of the web stro a bound together and very long, the 
longest reaching nearly ? of the whole ‘length of the web, 
like the str ings of a musical instrument. If one blows from 
the outer side upon the broad web it comes into vibration, and a 
sound is heard, which, though fainter, resembles very closely the 
well-known neigh.” He then fastened the outer or first feather 
with fine thread to a piece of steel wire and fixed it to a 4-foot 
stick, and found if he drew this with the outer edge of the feather 
through the air, at the same time making shaking motions of 
the arm to represent the shivering of the wings during flight, 
he was able to produce the neighirg sound with astonishing 
exactness. In bringing the matter before the Zoological Society 
Mr. Wolley confirmed Meves’s experiments. These experiments I 
hope to explain more fully anon. 

John Hancock, in 1875, in the ‘ Birds of Northumberland and 
Durham,’ vol. vi. pp. 105-113, severely criticises Meves’s theory 
and experiments. 

I cannot quote his article at length, for it is a very long one. 
He argues, from the diversity of structure exhibited by the rectrices 
of various species of Snipe, that they cannot be musical instruments. 
He failed to produce the neighing sound of a Snipe by Meves’s 
experiment, but admits “‘ when the web of almost any firm feather 
is blown upon a low vibrating sound is produced ; and such a sound 
is stronger when a tail-feather of the Common Snipe is used, 
arising apparently from the fact that the inner web is wide and firm ; 
but the sound is so low that it cannot be heard many yards off.” 

Later: ‘The sound is audible at a great distance, even when 
the bird has risen high into the air. No sound that could be 
produced under any circumstances by such feeble instruments as 
the lateral tail-feathers of the Snipe, instruments not larger than 
the wings of a Dragonfly, could be heard at any considerable 
distance. And it is scarcely to be doubted by anyone that the 
wings of a Snipe vibrating rapidly will produce some sound louder 
than any that could be made by a pair of small tail-feathers of a 
bird rushing downward through the air.” 

Prof. Altum (‘Ornithologisches Centralblatt,’ Oct. 1880) satisfied 
himself that he could produce the sound with the lateral tail- 
feathers. He, however, quotes “two adverse cases ” :— 

(1) A Snipe was observed “ bleating” as it sat, or rather stood, 

on an elevation. 

(2) A certain Alex Schmidt winged a Snipe which, with tail 
stiffly expanded, began to bleat in his hand, the air blowing 
through the web of the feathers, the wings being held 
close to the bird’s side. Every time the bird was moved 
rapidly against the wind his object was attained. 

Tt is to be noted that in both cases a strong wind was noticed 

to be blowing. ‘That these two cases are easily explicable I hope 
to adduce evidence later on. 

1907. | ‘‘BLEATING” OF THE SNIPE. 15 

In answer to this paper, von Zoppritz (Ornith. Centralblatt, 
Nov. 1880) disputes Altum’s tail-theory, giving four reasons, 
which JI will not here quote*. 

III. The wing-theory has had the greatest number of adherents. 

Macgillivray in 1840 (‘ British Birds,’ vol. iv. p. 372) expresses 
his conviction that this is the case. Sir Wm. Jardine, in the 
‘Naturalist’s Library’ (vol. xxvi. Ornithology, p. 180), says: ‘‘The 
sound is never heard, except m the downward flight, and when 
the wings are in rapid and quivering motion. ‘Their resistance 
to the air without doubt causes the noise. Dr. Saxby (‘ Birds of 
Shetland, p. 204) expresses his conviction that ‘ drumming is 
produced by the vibrations of the wings alone.’ ” 

Naumann, in the article already quoted, and Zoppritz (Ornith. 
Centralblatt, Nov. 1880) wrote in favour of it in Germany. 

John Hancock, whom I have already quoted, agrees in the 
main with what Mr. J. E. Harting (‘ Essays on Sport and Natural 
History,’ pp. 284 ef seq.) has written :—‘‘ From the peculiar 
vibration of the wings in the downward descent of the bird, it 
would appear that the primaries, instead of firmly overlapping 
each other, are, in the act of humming, turned broadside in the 
air, which is thus able to play across the inner web of each, and so 
to impart to each a vibratory motion and consequent sound—faint 
indeed in the case of a single feather, but audible enough when 
an entire wing is acted upon. Whether this be the true expla- 
nation of the singular sound, it is, of course, not easy to prove 
conclusively ; but it has certainly been accepted as such by many 
naturalists in England, who are the more inclined to adopt this 
view from having observed Peewits, Rooks, Gulls, and other birds, 
with tails very different from that of a Snipe, make an analogous 
sound while falling through the air.” That Mr. Harting was 
partially successful in producing the bleat artificially is evident, for 
he has ‘“‘ succeeded beyond expectation in producing a sound like 
the ‘ humming’ of the Snipe.” Again: “ But any of the primary 
wing-feathers will give forth a faint sound, which may be increased 
in proportion to the number of them passed through the air at 
once.” But he finds that the tail-feathers when fastened into a 
switch do not occupy the position they do naturally in the bird’s 
tail, because they are drawn through the air at right angles to the 
direction of flight, ‘‘in a position which is occupied naturally by 
the primaries, but unnaturally by the tail, and hence it must be 
the primaries (collectively) which produce the sound in nature. 
In this our sense of hearing is assisted by the sense of sight, for 
a perceptible vibration of the quill-feathers is observed every time 
the bird descends.” 

IV. Two observers maintain that the sound is produced by 
the agency of both the wing- and tail-feathers. 
H. Gadumer (‘ Naumannia,’ 1853, pp. 411-413) watched a bird 

* (Mr. F. W. Headley (° Nature’, vol. Ixx. p. 103, 1904) supported the theory that 
the drumming was produced by the outermost tail-feathers, and adduced an expe- 
riment by which the sound could be produced artificially. |—Ep. P. Z.S. 

16 MR. P. H. BAHR ON THE (Jan. 15, 

bleating, with a good field-glass. “The air pressing between the 
wings and the tail (a natural parachute) causes all the feathers of 
the tail and wings to vibrate and gives rise to the bleating sound 
—which is modified by stronger or weaker vibrations.” 

Text-fig. 5. 

a. Snipe bleating, showing characteristic position. 
b. Formation of tail as ordinarily held in flight. 

1907. ] ‘““BLEATING” OF THE SNIPE. 17 

Capt. W. V. Legge, in his appendix to the ‘ Birds of Ceylon,’ 
expresses the opinion that the sound is produced by the combined 
action of the wings and tail. 

I have dwelt on the literature for some length, in order that 
one may review the evidence adduced by the adherents of the 
different theories. 

In the summer of 1904, in the Fens of Cambridgeshire, I began 
to observe the Snipe in the act of bleating through a strong prism 
binocular, I had read none of the literature on the subject, and 
so had no preconceived ideas. The observations I made then I 
have had ample opportunities of confirming. 

I find that ordinarily the bird flies up to a height of 60-100 feet 
above ground, in windy weather going higher, with its tail held 
in the ordinary position of flight (text-fig. 3, 6), then, turning, it 
spreads its tail out like a fan, the two outer tail-feathers being 
spread out well in front of the other twelve and held firmly there 
(text-fig. 3,a). Immediately the bird begins to descend the bleat 
is heard (making due allowance for the time it takes for sound to 
travel). While descending the bird makes tremulous motions 
with its wings from the radio-carpal joint. The descent is made 
from 30-40 feet and occupies 2-3 secs., the bleat lasting the same 
time. The bird does not drop head foremost through space, but 
at an angle of from 45°-60° with the horizon. The tail as a whole 
is not vibrated, but it is quite easy to see the two outer tail-feathers 
with a strong glass vibrating to such an extent that their terminal 
portions become indistinguishable. Snipe begin to bleat in March, 
but if the weather is mild, in February, and continue to the end 
of May, though I heard one last year in Sutherland still bleating 
on June 25th. 

At the beginning of the breeding-season they may be seen 
bleating in pairs; but later on, when the hen is sitting, the cock 
bird may be seen performing alone over the marsh where the nest 
is placed. Under favourable conditions many bleat together, 
circling round the same spot for hours. On April 12th of last 
year, I had the good fortune to hear no less than twelve birds 
bleating together, a concert which they kept up all through the 
night. Every now and again, as if by common consent, there 
would be a lull, and all the birds would settle, but directly one 
began again ali the rest immediately joined in the chorus. 

Snipe bleat best in the early morning and in the evening, 
especially when the weather is dull and damp. It may be of 
interest to note that last spring I saw a specimen of the melanistic 
variety (Sabine’s Snipe) bleating. 

Once having convinced myself that the two outer tail-feathers 
are invariably spread out beyond the others, a fact which is now 
obvious to me with the unaided eye, it seemed to me that the two 
outer tail-feathers must be the active agents in causing the bleat. 
I accordingly procured several tails of the Common Snipe, and 
taking the two outer tail-feathers, pierced the shaft with a pin, 

Proc. Zoou. Soc.—1907, No. II, 2, 

18 MR. P. H. BAHR ON THE [Jan. 15, 

to which I firmly bound it with cotton and inserted the feathers 
into a cork at the end of a stick some six inches long. A hole is 
bored at the other end of the stick and a long string attached. 
This is whirled round the observer’s head and a typical bleat is 
produced. The second outer tail-feathers (sixth pair) produce a 
fainter sound, though this varies much in individual tails, the 
others make no sound at all. 

Text-fig. 4, 

Varieties of outer tail-fegthers of Gallinago celestis. (Natural size.) 

1907.] ‘““BLEATING” OF THE SNIPE. 19 

In order to ensure the success of the experiment it is necessary 
(1) that the feathers be placed so that the narrow edge, the outer 
web, shall encounter the resistance of the air; (2) that the feather 
be firmly bound to the pin, so that it cannot turn on its support ; 
(3) that the string be tied to one end of the stick, so that the 
long axis of the stick makes an angle with the direction of the 
string, if I may so put it, so that a vibratory motion is imparted 
to the stick as a whole, thus simulating the tremulous motion 
of the Snipe’s wings during the descent; (4) lastly, that the 
apparatus be moved at a uniform rate and not too fast. 

It is then found that after a period of silence the feathers begin 
to vibrate : first, the long-drawn-out note, which I may represent 
as “whti whtutu,” becomes gradually audible, it is then succeeded 
by a series of high and low notes ‘ bah-bah-ah-ah,” resembling 
the bleat of a young goat, lasting 3-5 secs., followed by a pause 
of equal length. This is repeated as long as the apparatus is 
revolving at a uniform rate. It is found that the individual tail- 
feathers, of which I collected a good number during the winter, 
vary considerably both in size, breadth, and markings, and, as 
might be expected, the note produced varies according to their 
physical characteristics. Thus a long narrow feather produces a 
sound of far higher pitch than a broader one of the same length 
(vide text-fig. 4). This fact I have noted when comparing the 
sound made by several birds when performing the nuptial evo- 
lutions over their breeding-grounds. To ascertain which part of 
the feather is essential in the production of the sound, I have cut 
off the narrow outer web, without altering the bleat in any way ; 
but if the barbs of the inner web be so disarranged that there is 
a break in their continuity, the web ceases to vibrate, and no 
sound is produced. That the vibration of the inner web is the 
active causative agent may be seen by the following simple 
experiments :—The feathers are attached to a cork, with the outer 
web held away from the observer, so that the narrow outer web 
shall cleave the resistance of the air. Thus affixed they are held 
out of the window of a train, or while riding a bicycle. As the 
resistance of the air is encountered the inner web begins to 
vibrate, slowly at first, but as the train gains speed, so rapidly 
that its outline is entirely lost, and it becomes a blurr; a low 
humming sound is at first heard, which soon reaches the typical 
pitch of the bleat. When the train has reached the speed of 
some 20 miles an hour, the whole feather will vibrate on the pin. 
If the feathers are at all loose on their pins it 1s curious to observe 
how they will always turn round so that the narrow outer edge 
encounters the resistance of the air. Furthermore, if the feathers 
be damped, they appear to act better, thus explaining, perhaps, 
why Snipe are found to be lable to bleat in damp weather. I 

-think this simple experiment readily explains away the ‘“ adverse 
eases ” of Prof. Altum (‘ Ornithologisches Centralblatt,’ Oct. 1880) 
already mentioned. 

That the bens bleat as well as the cocks is now, I suppose, a 
well-known fact (cf. von Preen, ‘ Naumannia,’ 1856, pp. 426, 427, 



20 MR. P. H. BAHR ON THE [Jan. 15, 

and Meveg, Proc. Zool. Soc. 1858, p. 200). I have observed it on 
several occasions myself. In the summer of 1902 I found four 
newly hatched Snipe in a patch inhabited by only a single pair; 
while lying concealed in the neighbourhood I observed repeatedly 
both old birds drumming above me. From the similarity of 

Text-fig. 5. 

a. Dorsal view of musculature of tail of Snipe. 
C=levator coccygis. 
- D=pygostyle. 
E=fused spines of caudal vertebre. 

A=slip of m. ilio-coccygeus to outer 
B=m. ilio-coccygeus. 

b. Ventral view of musculature of tail of Snipe. 

D=depressor coccygis. 

A=m. pubococcygeus ext. 

B=m. pubococcygeus. 
C=m. caud. ilio-femoralis. 

structure of the tail-feathers in both sexes, a fact which I have 
ascertained by dissection, one would infer that both sexes 
drummed. I cannot, however, agree with Meves that “as the 
feathers of the hen are generally less than those of the cock bird, 
the noise also made by them is not so deep as in the other case ” 

1907. | “‘ BLEATING” OF THE SNIPE. 21 

(op. cit. p. 200). I can find no difference either in the length of 
the feathers or in the intensity of the sound produced by the 
feathers of either sex. I have received a letter from Mr. 8. A. 
Buturlin, in which he says that in 1905, on the Kolyma Delta, he 
frequently observed both sexes of the eastern representative of 
our species (Gallinago raddit) drumming. 

Since the two outer feathers are extended beyond the other 
twelve during the descent, as I have described, I sought to find 
by dissection a mechanism by which this might be produced. On 
examining the tail of a freshly-killed bird, it is quite easy, by 
spreading out the tail, to make it assume the arrangement shown 
(text-fig. 3). I was unable, however, to find any special muscle 
peculiar to the species controlling the outer two tail-feathers. 
The muscle pubococcygeus ext. (text-fig. 5, 6) is inserted into the 
base of the shaft of the outer two tail-feathers, and is quite capable 
of performing this function. This muscle is to be found equally 
well developed in the other species of Plovers and Waders which 
IT examined. The nomenclature of the muscular system of the 
tail is that of Gadow in Bronn’s ‘ Thier- Reich.’ 

I have tried the same experiments as I have just described with 
the primaries from the wing of the Snipe, and was not able to 
produce any more sound with them than with others taken from 
other kinds of Waders, Pigeons, &e. There seems to have existed 
an opinion at one time that the bird produces two sounds, one 
with the wings and the other with the tail, the former being 
known as humming or drumming, and the latter whirring or 
bleating, produced while the bird is on the ground (cf. ‘ Zoologist,’ 
1881, p. 212, and 1846, p. 1501). I cannot say that this agrees 
with my own experiences. 

An Hxamination of the Structure of the Tail of Gallinago ccelestis 
(text-figs. 6, A, and 7). 

The normal number of feathers in the tail of this species is 14. 
It could hardly fail to strike the observer, on examining the tail, 
that the outer two differ considerably from the rest. Firstly, 
they are lighter in colour and their texture is firmer. On closer 
examination the shaft is seen to be strong and firm, presenting a 
decided outward curve towards its lower third. The outer web 
is narrow, and formed of stiff rami, which can easily be separated. 
The inner web, on the other hand, is extremely broad, being six 
times as broad as the outer, and formed of long stiff rami, of which 
some reach quite three-fourths the whole length of the feather, 
making a very acute angle at their insertion with the stem (text- 
fig. 6). The individual rami adhere firmly to one another, and 
can with difficulty be separated. These are provided with two 
well-developed rows of radii, the distal and the proximal rows 
(text-fig. 7, B), the former are twice the length of the latter. I 
must here express my great indebtedness to Mr. W. P. Pycraft, 
who has allowed me to make full use of his excellent paper, on 

Zo MR. P. H. BAHR ON THE [Jan. 15, 

“The Interlocking of the Barbs of Feathers” (‘ Natural Science,’ 
vol. ii, No. 19, Sept. 1893), and from which the illustration (text- 
fig. 7, A) is copied. Under the microscope the distal row is seen 

Text-fig. 6. 

A. Half of tail of Gallinago celestis from without inwards left to right. 
B. Half of tail of G. delicata from without inwards left to right. 
C. Half of tail of G. gallinula. 

to be well provided with hamuli and cilia (text-fig. 7, B). The 
hamuli deserve attention (text-fig. 7, C), since I believe them to 

1907. ] ‘“BLEATING” OF THE SNIPE. 23 

Text-fig. 7. 

> 3 3 
> ; \ ll 
MIE Wii) Y , ij py yp he Y) gr 




Wr hamuli 


A. Section of two rami of a feather showing interlocking of distal and proximal 
radii. (After W. P. Pycraft.) 

B. Ramus of Gallinago celestis, showing proximal and distal rows of radii. 

C. Distal radius of G. celestis. D. Proximal radius. 

K. Distal radius of middle tail-feathers of G. celestis. 

24. MR. P. H, BAHR ON THE [Jan. 15, 

be the essential factor in producing the bleat, in that they hold 
the stiff rami together like the strings of a harp. They are 
seven or eight in number, a number in excess of any other 
species of Snipe, and are well-formed, possessing a well-hooked 
terminal portion, which interlocks with the upturned edge of the 
radi of the proximal row (text-fig. 7, A & D). The outer web is 
formed of stiff rami, which possess rudimentary radii unprovided 
with hooklets. 

Of the remaining feathers, the sixth pair most nearly approaches 
our type feather in structure (text-fig. 6, A). The shaft is, however, 
not so strong, the outer web is broader, the inner narrower, and 
the rami are not so long, nor do they form such an acute angle 
with the stem (text-fig. 6,A). The hamuli are fivein number and 
not so well-formed, and, as I have said before, the sound produced 
by the vibration of the inner web of these feathers cannot compare 
in intensity with that produced by the outer pair. Thus the 
outer web becomes gradually broader, the inner gradually 
narrower as we reach the central tail-feathers (text-fig. 6, A), and 
the rami become progressively weaker, the hamuli fewer in 
number. Thus the distal radius of the middle tail-feathers 
possesses but four feebly curved hamuli (text-fig. 7, E). 

I have examined the tail of this species during the moult. On 
the 17th of August, 1906, I received several from Scotland, just 
at the beginning of the moult. The outer tail-feathers, I find, 
have lost much of their bleating power and the note produced is 
not so intense. On microscopical examination I find the cilia 
(text-fig. 7, C) have all been worn away. From this I infer that 
the cilia play a certain part in the production of the sound. 
From the 17th August to the 6th September I received 
many tails in which the new feathers were just growing, and I 
find that in every case the outer tail-feather (the sonorous instru- 
ment) is the last to be assumed. The newly assumed feather 
possesses fu]l bleating powers. I have also examined feathers 
from young birds of the year directly they have assumed their 
full plumage; these, I find, will bleat as well as those of a fully 
adult bird, and possess the normal structure and characteristic 
number of hamuli. 

One variety of G. celestis still remains to be mentioned, i. e. 
Gallinago raddw (Buturlin), which is the eastern representative 
of our species and is much lighter in colour. With characteristic 
kindness, Mr. Buturlin has sent me skins of this and several other 
species from the Kolyma Delta in Siberia (69° 4’ 20” N. and 
160° 55" E.), accompanied by most valuable notes, for which I am 
deeply indebted. The feathers of the tail, of which I have figured 
a specimen (text-fig. 4, 4), behave in the same way as those of our 

Examination of the Tail of other Species of Gallinago. 

Gallinago delicata, or Wilson’s Snipe, of N. America, differs 
in the eyes of some materially from G. c@lesiis in possessing 


sixteen tail-feathers. Of these the outer two are specialised, but 
differ from those of G. celestis in that the mner web of these 
feathers instead of being broader is narrower than that of the 
other (text-fig. 6, B); they are, in fact, somewhat attenuated—a 
process which, as we shall see later, reaches its extreme in the 
Pin-tailed Snipe (G’. stenwra) of India. The shaft is strong, but 
not so strong as that of our species. The inner web is three 
times as broad as the outer. Both feathers will produce a 
bleat on experiment; the sound is of a far higher pitch than 
that of G. calestis, as might be inferred from the character of 
the feathers, and is what is aptly described by the Americans 
as “winnowing.” The rami are shorter in comparison with 
G. celestis, and make an acute angle with the shaft. The radii 
of the outer web are rudimentary, of the inner web the distal 
is but one-third longer than the proximal row; the hamult 
are five in number, but are not so well hooked as those of 
G. celestis. Of the two, the outer or eighth is more attenuated 
than the seventh pair. 

Regarding the habits of this species I have the following 
references :— 

Baird, Brewer, and Ridgway, ‘ Water-Birds of N. America, 
vol. i. p. 191 :—‘“‘ Capt. Blakiston noticed that this species per- 
formed the same evolutions as the European bird, this usually 
about sunset, but at times continuing 13 hours later. The noise 
made on these occasions he compares to rapidly repeated switches 
of a cane in the air, and this was repeated every half minute with 
cecasional longer intervals. The sound lasted about three seconds, 
and was made as the bird descended rapidly in a vertical direction, 
being caused apparently by the quill-feathers of the wings. This 
sometimes took place in the middle of the day, but only during 
the love season.” 

Again :— 

Audubon, ‘ Birds of America,’ p. 343 :—‘ These birds are often 
met with in meadows, or on low grounds, and by being on the 
spot before sunrise, you may see both mount high in the air in a 
spiral manner, now with continuous beats of the wings, now in 
short sailings, until more than a hundred yards high, when they 
whirl round each other with extreme velocity and dance as it 
were to their own music, for at this juncture, during the space of 
5 or 6 minutes, you hear trolling notes mingling together, each 
more or less distinct, perhaps according to the state of the 
atmosphere. The sounds produced are extremely pleasing, though 
they fall faintly on the ear, but I am well assured that they are 
not produced simply by the beatings of the wings, as at this time 
the wings are not flapped, but are used in sailing swiftly in a 
circle not many feet in diameter. A person might cause a sound 
somewhat similar by blowing rapidly alternately from one end 
to another across a set of small pipes consisting of 2 or 3 

From this we gather that this species performs its evolutions at 

26 MR. P. H. BAHR ON THE [Jan. 15, 

a greater elevation than our species, also that Audubon noticed 
the fact that both cock and hen bleat. 

Wilham Brewster, in Chapman’s ‘ Handbook of Birds of North 
America,’ writes, pp. 154-155 :—‘‘In the springtime, and occa- 
sionally in autumn also, Wilson’s Snipe mounts to a considerable 
height above his favourite meadows, and darts downward with 
great velocity, making at each descent a low yet penetrating 
tremulous sound, which suggests the winnowing of a domestic 
Pigeon’s wings, and if heard at a distance, the bleating of a goat, 
and which is thought to be produced by the rushing of the air 
through the wings of the Snipe. This performance may be 
sometimes witnessed in broad daylight, when the weather is 
stormy, but ordinarily it is reserved for the morning or evening 
twilight or for moonlight nights, when it is often kept up for 
hours In succession.” 

Other American species deserve mention here :— 

Gallinago nobilis (text-fig. 8, B) has 16 tail-feathers, of which 
the outer three are attenuated and the fourth partially so. It 
inhabits Ecuador and Colombia, and is allied to G. australis, 
which species I shall treat of later. I can find no reference to the 
breeding-habits of this species. On experiment the three outer 
feathers bleat well. As in G. delicata the rami of the outer web 
possess but rudimentary radii; those of the inner, however, possess 
a distal row which is one-third longer than the proximal, and the 
former is provided with five hamuli which are not well-hooked. 
The rami are thicker and stiffer than in the aforementioned species, 
and I suspect it is more by the vibration of the rami as a whole 
that the sound is produced, in contradistinction to the vibration of 
the inner web alone as in @. celestis and delicata. The inner web 
of the eighth pair is but little broader than the outer, but that of 
the seventh pair is nearly twice as broad, thus resembling the 
outer tail-feathers of G. delicata. The middle tail-feathers conform 
to the ordinary type. Thesound produced is hard to describe; it is 
flute-like, but possesses a definite bleating character. 

Gallinago frenata.— Another South American species inhabiting 
Brazil, believed to be a Neotropical form of G. delicata. This 
species has 16 tail-feathers, of which the outer four are 
attenuated, the outermost being one-fifth inch in diameter, the 
inner being but slightly wider than the outer web. The inner web 
becomes progressively larger towards the centre of the tail. . This 
Species agrees with the foregoing in having the specialised 
feathers of a lighter colour: all four bleat; this is similar to that 
of G. nobilis, but shriller: microscopically they resemble the 
structure of G. delicata. The distal radii of the inner web are one- 
third longer than the proximal row and possess five well-curved 
hamuli. The rami of the outer web are stiff and structureless. 

G. paraguay is a subspecies of G. frenata inhabiting Paraguay. 
It is much larger than the type, and the tail-feathers are 

1907.] ‘‘BLEATING” OF THE SNIPE. 27. 

consequently larger and the bleat deeper in tone ; otherwise they 
agree in number and structure. A description of the breeding- 

Text-fig. 8. 

A. Half of tail of @allinago major from without inwards left to right. 
B. Half of tail of G. nobilis from without inwards left to right. 

C. Half of tail of G. stenwra. 

habits of this species is to be found in a paper by Durnford in the 
‘This, 1877, p. 198:—‘ During the spring they go through the 

28 MR. P. H. BAHR ON THE Jan. 15 

same aerial movements as the Common Snipe at home, rising to 
a great height by a circling motion, and ‘drumming’ whilst 
descending in a diagonal line. How is this curious habit to be 
accounted for in the South-American and European forms, except 
by the theory of inheritance from a common progenitor ?” 

Gallinago australis —Latham’s Snipe has 18 tail-feathers, of 
which the outer three are attenuated, two being less than 3 of an 
inch in diameter; the outer six, however, are doubtlessly specialised, 
as they differ markedly from feathers from the centre of the tail 
both in structure and colour. The feathers bear a certain resem- 
blance to those of the South-American species. The shaft is 
thick and curved ; the rami of the inner web are long and thick, 
but more easily separated than in the latter species. The rami 
are peculiar, in that they are thicker and stiffer than in any other 
species. The distal rami of the inner web are longer than 
the proximal row and are provided with 5 hamuli. The rami of 
the outer web are stiffand structureless, thus resembling G. celestis, 
paraguaye, and frenata. The feathers produce aloud bleat, some- 
what similar to that of G. celestis. 

I have received from Mr, Alan Owston, of Yokohama, a skin of 
this species, accompanied by some very valuable notes, which 
add materially to our knowledge of the habits of this species, for 
which Iam greatly indebted to that gentleman. He says: ‘“ They 
breed on the grassy moorland at the foot of Mt. Fugiyama, at an 
elevation of 2000-3000 ft. above the sea (Fugiyama is 12,500 ft. 
high). I have watched them on the 28th April, and on other 
dates during the breeding-season. When alarmed they fly round 
overhead, circling round generally against the sun, and every now 
and again they begin to cry ‘chip, chip, chip, sheep, cheo, che- 
cheo,’ and then rush downwards at the intruder, beating the air 
in the descent and making a terrific rushing noise.” 

He also sends me an extract from Capt. T. W. Blakiston’s 
notes on the breeding-habits of this species published in the 
‘Chrysanthemum’ for Nov. 1882, p. 524 et seg., referring to 
“Birds observed on the 8.E. coast of Yezo in May” :—“ The 
Australian species act very like the Snipe of North America, 
by flymg round pretty high and making sudden rapid descents 
almost to the ground, which latter movement is accompanied by a 
whisping noise. At evening and during the day in dull weather, 
these evolutions are commonly performed; and in dirty rainy 
weather the noise is heard even in the middle of the night.” 

Gallinago aucklandica.—Resembles G. australis in having 18 
tail-feathers, the outer three of which are attenuated; they are, 
however, much softer in structure than in that species. The rami 
of the inner web are easily separated, and possess 4 hamuli; those 
of the outer are provided with rudimentary rows of radii, thus 
approximating to certain Asiatic members of the genus. I have 
received a specimen of this rare species from the Christchurch 
Museum, N.Z. No mention of any bleating-habits is made in 

1907.] ‘“‘BLEATING” OF THE SNIPE. 29 

the literature. Contrary to what one would expect from its 
peculiar Rail-like appearance, the tail-feathers produce a very 
distinct and pleasing sound of a high-pitched character, some- 
what resembling that of certain Asiatic species. 

Gallinago equatorialis (nigripennis) is an inhabitant of Central 
Africa, south and east of the great desert. It possesses 14 tail- 
feathers, of which the outer four are attenuated, and are less than 
1 inch in diameter ; they are pure white. This form is nearly allied 
to the Common Snipe. I have been quite unable to procure either 
any feathers or a skin of this species. 

Its habits are described in Reichenow’s ‘ Végel Afrikas,’ Band i. 
p- 236 :—“ This bird is called Spook Vogel by the Boers, on account 
of its drumming cry, which the bird makes in the morning and 
evening during its flight.” 

Gallinago gallinula.—The Jack Snipe has 12 tail-feathers, of 
which the outer three are markedly shorter than the three central 
ones (text-fig. 6, C). 

Their texture is soft and the rami are easily separated, in 
contradistinction to those of the species we have already con- 
sidered. On experiment these feathers produced no sound at all. 

The structure of the outer web of the outer feathers more 
nearly approaches that of the inner—a marked difference to that 
found in the other feathers we have been considering ; that is, the 
rami of the outer web are provided with distal and proximal rows 
of radii and thus adhere together. The distal radii are provided 
with 4 hamuli both in the outer and inner webs. 

The breeding-habits of this species were described originally by 
Wolley in Hewitson’s ‘Eggs of British Birds,’ vol. ii. p. 356 :— 
“Tt was on the 17th June, 1853, in the great marsh of Muonio- 
niska (Finland), that I first heard the Jack Snipe, though at that 
time I could not guess what it was—an extraordinary sound, unlike 
anything I had heard before ; I could not tell from which direction 
it came, and it filled me with a curious surprise. My Finnish 
interpreter thought it was a Capercally, and at that time I could 
not contradict him. I know not better to describe the noise than 
by likening it to the cantering of a horse in the distance, over a 
hard hollow road, it came in fours with a cadence, a clear yet 
hollow sound : it was not long afterwards that I ascertained the 
remarkable hammering noise in the air was made by the Jack 

Mr. 8. A. Buturlin, in sending me a specimen of this species, 
with characteristic kindness writes the following notes of its habits 
as observed on the Kolyma Delta :—“ Its drumming is exceedingly 
like the noise of a cantering horse on a hard road, as so well 
described by one of the best field-observers—the late John Wolley. 
I heard it every day in the summer of 1905, when on the 
Kolyma. The bird usually flies so high, that even with the aid 
of the midnight sun and good Zeiss binoculars it is often quite 

30 MR. P. H. BAHR ON THE [Jan. 15, 

invisible; nevertheless the sound ‘ top-toppy-top-toppy * is quite 
clearly heard.” 

I might also mention here that the remarkable sound produced 
by the W ood-Sandpiper (7'otanws glareola) was found by Mr. Buturlin 
to be vocal. He shot a score of them ‘“ drumming ” while sitting 
on a branch of Ulmus incana, or some local Salix, on the Kolyma, 
and has observed them for a long time at the distance of a 
few yards. The male flies about in wide circles, beating his wings, 
now floating on outstretched wings uttering as loud but not such 
hollow notes as the Jack Snipe. 

I can only say at present that, in view of the failure to produce 
the drumming of the Jack Snipe artificially, I suspect there must 
be some other mechanism by which the sound is produced. 

Gallinago major: “'The Great Snipe” (text-fig. 8, A).—This 
species has 16 tail-feathers, of which the outer four are white. They 
are somewhat shorter than the feathers from the centre of the tail, 
which are similar in all species of Gallinago. The feathers produce 
no sound on experiment. The rami are soft, like those of G. gal- 
linula, and can easily be separated. The outer web is composed of 
rami provided, as in the case of G@. gallinula, with rows of distal 
and proximal radii, of which many are well developed. In the 
inner web the distal row is one-third longer than the proximal, 
and is provided with 4 feeble hamuli. The rami are inserted into 
the shaft at an obtuse angle. 

Its breeding-habits have been described by Prof. Collett, of 
Christiania, in Dresser’s ‘Birds of Europe,’ vol. vil. p. 635 :— 
“The Double Snipe is chiefly a nocturnal bird. Not only does it 
migrate at night, but it is in motion almost solely after twilight, 
when its peculiar ‘spil* or drumming takes place; and it also 
searches after food chiefly during this time of the evening. ... 
Tt has a so-called ‘ Leg’ or ‘ Spil,’ like some of the Grouse tribe, a 
sort of meeting-place, where they collect to ‘drum’ and often to 
engage in combat for the possession of the females. . . . It does not 
indulge in aerial evolutions, but remains on the ground. ... The 
male bird utters a soft, almost warbling note, which is accompanied 
by a peculiar snapping sound caused by striking the mandibles 
together several times in quick succession. If a person approaches 
one of these drumming-places he can hear at some distance the 
low note: ‘bip bip, bipbip, bipbiperere, biperere’; and when 
within 100 paces, if the night is still, he begins to hear other 
peculiar sounds. ... Whilst producing these notes the bird is in 
ecstasy and raises and spreads its tail like a fan, the outer tail- 
feathers showing in the halt-darkness like two white patches.” 

_ Here, again, having no experience of the aforementioned habits 
myself, I can only conjecture that the sound is vocal. 

Now we come toa group of Asiatic species, which bear much 
resemblance to each other in the structure of their tail-feathers :— 

Gallinago solitaria (text-fig. 9, «).—According to the formula 
for this species given in Seebohm’s ‘ Geographical Distribution of 

1907. | ‘““BLEATING” OF THE SNIPE. 31 

the Charadriide,’ this species ought to have 18 tail-feathers ; 
but a specimen sent me by Mr. Buturlin from the Western Tian 
Shan Mountains has 20 tail-feathers. The outer six are attenuated, 

a. Tail of Gallinago solitaria. | b. Tail of G. megala. 

the first five markedly so. In this case both the outer and inner 
webs are very much narrower. Unlike any of the preceding 
species, the rami of the outer web are provided with fully developed 

32 MR. P. H. BAHR ON THE [Jan. 15, 

vows of distal and proximal radii, the former being provided with 
4 hamuli in the same manner as the inner web. Herein this 
group differs notably from any of the preceding; the rami are 
short and stout, proximal and distal rows of radii the same size. 

The bleat produced by these feathers is very loud, and consists 
of a number of notes of different pitch intermingled, caused 
apparently by the difference in breadth of the different musical 
feathers in the tail. 

T venture to think that in this group the bleat is produced by 
the vibration of the feather as a whole, not by the inner web 
alone as in G. calestis. 

An account of its breeding-habits is to be found in Hume and 
Marshall's ‘Game Birds of India’ :—‘“ They are to be heard and 
seen in the higher portion of the hills, soaring to a considerable 
height, repeatedly uttering a loud sharp, jerky call, and then 
descending rapidly with quivering wings and outspread tail, pro- 
ducing a hard buzzing sound, something like, but shriller and 
louder than, that produced by G. celestis, though they do not 
descend as rapidly as the latter.” 

Gallinago megala.—This species inhabits 8.E. Siberia from 
Lake Baikal to the North Island of Japan. It possesses 20 tail- 
feathers, of which the outermost are 3 inch in diameter. The 
attenuated feathers are shorter than those from the middle of 
the tail (text-fig. 9, ). Microscopically they resemble those of 
G. solitaria in every way. The bleat they produce also resembles 
that of the foregoing species, but is far higher in tone. An 
account of its breeding-habits is to be found in Taczanowski’s 
‘Fauna ornithologique de la Sibérie orientale,’ p. 958, a refer- 
ence kindly given me by Prof. Newton. I have translated the 
passage from the French: quoting from Prjevalski he says :— 
‘That it retires to breed in the deepest marshes, covered with 
black scrub, and performs aerial evolutions as follows. The male 
soars up in the same way as our Snipe does, and after having 
described large circles in its flight above the place in which the 
female is nesting, it darts downwards in an oblique direction, 
making (probably with the rectrices, as does our Snipe) a loud 
sound, like the noise produced by a racquet when the handle has 
been broken. ‘This noise gains more and more in intensity as it 
approaches the ground, and ceases about 100 paces from it, and 
then the bird continues its flight, repeating a note, which one can 
express by tic-tic-tic.” 

T have received a tail of this species from Central Siberia from 
Mr. Buturlin, who sends me this account from his first part of 
the ‘ Limicole of Russia’ (1902, pp. 77-78), an account contributed 
by the late M. Schwedow :—‘ After 6 o’clock p.m. (in May, near 
Irkutsk) one can see in the woods or on forest-swamps the ‘forest 
Snipes’ wheeling round and round in the air, and nearly always 
repeating in rapid succession notes like ‘chwi, ehwi, chwi,’ or 
‘ zswee, zwee, zwee.’ From time to time one or other of the birds 
stops beating its wings, somewhat partially closes them, and 
swoops obliquely down, while vibrating notes are heard in the 

1907. | ‘“‘BLEATING” OF THE SNIPE. 33 

air, somewhat like water pouring in the distance, gradually 
becoming higher and higher, and more sharply falling on the ear. 
The bird falls like an arrow, but when some fathoms from the 
tops of the trees it suddenly stops in the air and at the same 
moment uttering ‘chic-ka-chee,’ flies on again. After making 
some rounds in the air the bird silently or with the same sound 
‘zswi, zwsi, goes higher and higher in the air and recommences 
the same performance.” 

Gallinago stenura (text-fig. 8, C).—The Pin-tailed Snipe of 
Tndia breeds in Hast Siberia, and possesses the greatest number of 
tail-feathers of the genus—26-28, of which the outer eight or nine 
are attenuated. The outer ones are so much attenuated that they 
actually do resemble pins. The outer feather measures 545 inch 
in diameter, the eighth ;, inch. 

The outer web resembles that in G. megala in possessing fully 
developed radii; the number of the hamuli in the first pair is 5, in 
the eighth pair 4. The ramiare short and thick. These feathers 
produce no sound on experiment. The passage from Swinhoe on 
Formosan Ornithology, ‘ Ibis, 1863, p. 415, is the quotation from 
Taczanowski which I have just given and refers to the preceding 

Mr. Buturlin writes of this species on the Kolyma :—‘ It 
was only one day, 25th June, 1905, that I could observe its 
breeding habits.... I watched it during two or three hours 
with strong binoculars at a distance sometimes of not more than 
200 yards.... It flew about uttering a high, loud, somewhat 
harsh note, not clear enough to be styled a whistle, like ‘ psait, 
psait, psait,’ and seemingly produced not by some mechanical 
means, but by its voice. From time to time (but not so often as 
our Common Snipe) the bird makes head foremost a dive in the 
air, just as our Common Snipe, but the descent is in time and 
distance quite twice as long as in the common species. When 
falling through the air the bird repeats its note more and more 
swiftly. ... At the lowest point of its descent, the bird holds the 
wings high over its back, just like a swiftly descending pigeon or 
duck, and then ascends several feet without evident motion of its 
wings. I could not see any opening or spreading out of its tail, 
when swooping downwards.” 

John Hancock, writing in his ‘ Birds of Northumberland and 
Durham,’ found great difficulty in accepting the tail theory, 
because of the diversity in structure of the tail-feathers to be met 
with in this genus, especially the feathers of G. stenwra, which, he 
says, are considered to be musical instruments. Thus he raises 
a very reasonable objection, which I shall here quote :—‘ Other 
species have the same almost webless feathers at the sides of the 
tail, varying only in number. Here then we see a species in 
which the so-called sonorous or ‘ musical’ feathers do not possess 
the structure, firmness of web, and length of rays, which appear 
to be mainly relied on as the sound-producers; though the 
rigidity and form of the shaft are in some way or other apparently 

Proc. Zoot. Soc.—1907, No. III. 3 


thought to have some influence in the production of the sound, 
independently of the rays or web. Were these feathers sonorous 
instruments, we should expect to find a greater uniformity in 
their structure. But, in fact, the tail-feathers of the true Snipes 
are remarkable for their diversity, so much so that the birds have 
been divided into four groups, and this mainly on account of a 
difference in the number and form of these feathers.” 

T have tried to show that the mechanism differs considerably in 
different species, just as the sound varies. 

For reasons stated before, I believe that the bleat of G. ccelestis 
is produced by: the vibration of the inner web as a whole; in 
the case of G. frenata, nobilis, and australis by vibrations of the 
individual rami; while G. megala and solitaria produce sounds of an 
entirely different character by vibration of the feather as a whole. 

Finally, I will but briefly mention two species belonging to 
closely allied genera :-— 

Scolopax rusticola.—The Wopdcock is known to perform certain 
evolutions during the breeding-season, producing at the same 
time a curious sound, which is acknowledged to be vocal. 
Certainly there is no evidence from examining the twelve feathers 
of the tail that any of them are specialised structures. The outer 
ones do not differ materially either in size or structure from any 
of the others. Microscopically the outer web is composed of plain 
rami provided with but rudimentary radii. The hamuli are four 
in number and their terminal portion is badly hooked. 

Philohela minor (Gmel.)—The American Woodcock has 14 
tail-feathers, of which the outer ones are decidedly shorter than 
the others; they produce no sound in experiment and are in 
macroscopical and microscopical structure similar to the last-named 
species. A good account of the habits of this bird in the breeding- 
season is to be found ih Chapman’s ‘ Birds of Eastern North 
America, p. 153 :—-“‘ He begins on the ground with a formal, 
periodic, peent peent, an incongruous preparation for the wild 
rush that follows. It is repeated several times before he springs 
from the ground, and on whistling wings sweeps out the first loop 
of a spiral, which may take him 300 feet from the ground. Faster 
and faster he goes, louder and shriller sounds his wing song; then, 
after a moment’s pause, with darting headlong flight, he pitches 
in zigzags to the earth, uttering as he falls a clear, twittering 
whistle. He generally returns to near the place from which he 
arose, and the peent is at once resumed as a preliminary to another 
round in the sky.” 

Certain of the primaries of the wing of this species are 
characteristically attenuated, for what purpose I am unable to 
discover, as they certainly do not produce any sound by any 
means I have employed. 

T do trust that in the enquiries I have made I may be followed by 


others. I cannot attempt to explain many of the facts I have set 
forth in this memoir; and yet an explanation ought to be forth- 
coming, and particularly in reference to the microscopical part of 
the subject on which I have mainly been able to dwell. The exact 
relations of the number and size of the barbules and hamuli to 
the sounds they produce is worth investigating, and still more 
is the cause of the breaks in the continuity of the sounds which 
you have heard—or, rather, not heard. This last would need the 
application of one who is intimately acquainted with the science 
of acoustics, which I make no pretence to be, and therefore I 
cannot offer any suggestion which will account for the non- 
continuity of the “ bleating” or ‘ humming”—its sudden stops 
and its sudden recurrence. There is much more to be learnt in 
this matter, and I would pray those who may be unconvinced by 
my experiments, at least to try to account for those marvellous 
sounds in some manner more satisfactory, and I assure them that 
there is no one who would be better pleased than myself to find 
that they can be so accounted for. 

In conclusion, I should lke to tender my sincere thanks to 
Prof. Newton, of Cambridge, without whose assistance the above 
facts would never have been recorded. 

Since reading this paper I have received a skin of a female 
specimen of G. equatorialis. The sound produced is disappoint- 
ing in volume; in tone it bears a resemblance to that of the bleat 

of G. calestis. 

3. Contributions to the Knowledge of the Systematic 
Arrangement and Anatomy of certain Genera and 
Species of Squamata. By Frank HE. Bepparp, M.A., 
F.R.S., Prosector to the Society. 

[Received December 7, 1906. | 
(Text-figures 10-19.) 

(1) On some Specific Characters of Chameleons shown in the Internal Organs, 

p. 3d. 

(2) Some Notes upon Chameleolis, p. 45. 

(3) The Position of the Umbilicus in certain Vipers, p. 50. 

(4) Some Notes upon the Anatomy of Zonwrus, with Special Reference to the 
Hyoid, p. 52. 

(5) Some new Facts bearing upon the Affinities of Gerrhonotus, p. 56. 

(6) On a Point of Structural Resemblance between Heloderma and Varanus, 
and on some Specific Characters of Varanus, p. 59. 

(1) On some Specific Characters of Chameleons shown in the 
Internal Organs. 

The external differences among Chameleons are plainly set 

forth in vol. iii. of Boulenger’s ‘Catalogue of Lizards in the 


British Museum.’ The differences between the viscera of different 
species are by no means so well known. Indeed the only recent 
memoir known to me dealing with the visceral anatomy of 
Chameleons, which also refers to specific differences within the 
genus, is that by Dr. Wiedersheim*, chiefly dealing with the 
respirator y system in Oh. vulgaris and Ch. monachus. 1 have 
dissected, with reference to more than one point in the visceral 
anatomy, the following species, viz.:—Ch. vulgaris, Ch. calcarifer, 
Ch. dilepis, Ch. pumilus, Ch. parvilobus, Ch. teniobronchus, Ch. 
basiliscus, and Ch. verrucosus. Iam in consequence able to offer 
some additional anatomical facts concerning the genus, which are 
also of classificatory importance. 

T may first of all call attention to an external character of the 
little-known Chameleon calcarifer. One of the external characters 
which distinguishes Ch. calearifer from Ch. vulgaris is the presence 
in the former of a more distinct ventral crest composed of a 
line of conical and, at times, overlapping “enlarged granules.” 
This line is traceable, but is by no means so well marked, in 
Ch. vulgaris. The division of these ventral scutes which marks 
the position of the umbilicus in the foetus is therefore exceedingly 
obvious in Ch. calcarifer and less easily to be mapped out in 
Ch. vulgaris. It lies behind the middle line of the body. It is 
represented by a long space contained in the middle of the ventral 
crest, which bifurcates to embrace it, and in this region therefore 
1s double. The number of scales on the two sides is uneven ; 
T counted 13 on the left and 11 on the right side. The size of the 
region of the integument which appears to mark the umbilicus was 
rather greater in “Ch. vulgaris, but, as already said, the indistinct- 
ness of the ventral crest renders ‘it difficult to be accurate. In 
Oh. dilepis this area was quite as distinct as in Ch. calcarifer and 
occupied the same position; there were, however, only 10 pairs of 
scales, closely apposed. 

The lungs show some variation in structure from species to 
species. That there is some variation in these organs has already 
been pointed out by Wiedersheim, who figures those of Ch. vulgaris 
and of Ch. monachus. The differences seem mainly to affect the 
number and form of the tags which are appended to the lungs, 
those very characteristic anangious outgrowths of the lung. 
Wiedersheim distinguishes between the more or less cylindrical 
outgrowths and the branches of the lung itself which bear them. 
The outgrowths, which exist more anteriorly and always on the 
ventral side of the lung, are simply the tubular ceca. Towards 
the end of the lung the lung itself is divided into several 
processes. In a young example the author quoted found that the 
ceca were solid. There is therefore reason for distinguishing the 
pulmonary ceca from the lungs, and a careful examination of both 
shows a ceasing of the reticular bands which cover even the 
anangious part of the lungs. As the number of outgrowths not 

* Ber. naturf. Ges. Freiburg-i.-Br., Bd. i. 1886, Heft 3. 


only varies in individuals, but also on the two sides of the body 
of the same specimen, their mere number and arrangement can 
hardly be utilised as distinctive of species without examining a 
large series. Yet I am disposed to think, for reasons that will be 
discussed later, that Ch. monachus does differ from Ch. vulgaris. | 

In Chameleon calcarifer the langs show the same general 
structure as do those of Ch. vulgaris. That is to say, the lung 
itself is frayed out into processes posteriorly. These again muy 
or may not give rise to the tubular cecal outgrowths. The latter 
show no network upon their surface, but the direction of the fibres 
of which they are partly composed is rather circular. On the 
ventral side also, some way in front of the end of the lung, the 
lung itself is prolonged into processes. I counted altogether in 
one lung examined fifteen tubular cecal outgrowths. But as the 
numbers have been stated by Wiedersheim to vary in Ch. vulgaris, 
the exact number is probably not a matter of importance. They 
were, however, certainly more numerous than in an example of 
Oh. vulgaris which I have myself studied. What appears to be 
of importance is to note that the lung itself is divided and that 
the tubular outgrowths do not arise from a ling with an entire 
margin. The subdivisions of the cavity of the lung seem to be 
exactly as Wiedersheim has described for Ch. vulgaris. In 
Ch. verrucosus the tubular cecal outgrowths are very numerous. 
T counted twenty-five or more of them. All the cecal outgrowths 
were borne in four tufts, of which that furthest from the bronchus 
was the largest, and consisted also of an outgrowth of the lung 
itself. The individual tubular ceca were frequently to be seen 
arising by the division of a common stem (text-fig. 10, p. 38). 
The disposition of the ceca im this species is very different from 
that which I have observed in others. 

The lungs of Chameleon dilepis appear to differ in certain respects 
from those of the species that have been hitherto described. The 
obvious difference is the tubular character of the cecal outgrowths, 
which have hardly any dilated termination, shown so plainly in 
Ch. parvilobus, for example (text-fig. 12, p. 39). In the second 
place, the tubular ceeca are thick-walled and not at all transparent 
except in parts, and then not so transparent as in other species. 
Furthermore, these processes are distinctly shorter in Ch. dilepis 
than they are in Ch. parvilobus, as 1s indicated in the annexed figure 
(text-fig. 11, p. 39). The differences above set forth can hardly be 
due merely to a different state of contraction, since both specimens 
came out of the same bottle of aleohol in which they had been 
preserved some time since ; and very well preserved, for there was 
no trace of softening or disintegration of the viscera. The ceca 
of Oh. dilepis are certainly to some extent contracted, as they can 
be pulled out without using undue force. There remains, however, 
a condition which differs from the attenuated and thin ceca of 

Ch. parvilobus and Ch. verrucosus on the one hand, and from the 
prolongation of lung-substance with shorter ceca in Ch. calearifer, 
‘on the other. The marked distinctness of the czeca from the lung 


is, in fact, a feature of this species as contrasted with those that 
have been mentioned. It will be observed in the figure (text-fig. 11) 
that the lung terminates in a cecum which continues in the same 
straight line. This seems to be the case also with other species. It 
suggests, of course, the distal terminal air-sac (abdominal air-sac) 

Text-fig. 10. 

Lung of Chameleon verrucosus, entire. 

of the bird’s lung. The arrangement of the other ceca is shown 
in the figure referred to. They are developed along a considerable 
region of the ventral margin of the lung. The larger number of 
the ceca are, however, massed at one spot, which is not at the end 
of the lung as in Oh. parvilobus, but at about its middle. Another 


noteworthy difference about these ceca as compared with those 
of other species which I have examined, is that there is an anasto- 
mosis between the roots emerging separately from the lung. 
Finally, the small number of czeca as compared, for example, with 
Ch. calearifer is a fact worthy of attention, since it is the beginning 
of the immense reduction seen in Ch. basiliseus, which culminates 
in the total absence of these ceca in Ch. pumilus. 

Text-fig. 11. Text-fig. 12. 

Text-fig. 11.—Lung of Chameleon dilepis, entire. 
Text-fig. 12.—Lung of Chameleon parvilobus, opened longitudinally. 

The above description also applies in generalities to the right 
lung of the same individual. That is to say, with regard to the 
shortness, tubular character, and fewness of the cecal outgrowths. 


A specimen which I had the opportunity of examining fresh 
showed how the lungs may vary in individuals, as was pointed 
out by Wieder sheim, This variation consists principally in the 
Jarger number of ceca. I should mention, however, first of all 
that there is just a shadow of a doubt as to the identity of the 
species. In the second smaller specimen the occipital lobes 
characteristic of the species were disproportionately smaller than 
in the larger individual, whose lungs I have already described. 
In the second .place, the larger individual had no trace whatever 
of the ‘‘ Hohlenvenenfortsatz” of the right lobe of the liver 
accompanying the postcaval vein. In the smaller individual there 
was a considerable process of hepatic tissue accompanying the 
postcaval vein for some distance. 

In the case of both right and left lung, the lung ended, in the 
same way as in the example already described, in one bifid caecum, 
bifid from the very first. In one lung I counted 14 other cecal 
outgrowths, of which five were particularly short. In the other 
lung I'found as many as 16 outgrowths, of which only three or 
four were short. Two, or even three, sometimes borne upon the 
same stem. The ceca are of considerable diameter and clubbed 
in form ; they contrast markedly with those of Ch. verrucosus. 

I have also selected for figuring the lung of Ch. parvilobus (text- 
fig. 12), which is at the very opposite extr emity from the other 
species figured in the present communication, viz. Ch. dilepis. The 
ceca are numerous and extremely slender and in some cases of great 
length. Thus the longest measures 33 mm. as against 32 mm. of 
the length of the lung itself, and there are several other czca 
nearly or quite as long. The longest of the diverticula are at the 
posterior end of the lung. The whole ventral border is also beset 
with diverticula, but these are invariably short; all show a marked 
dilatation at the free extremity. In contrasting these slender ex-. 
tended diverticula with the short thick diverticula of the two species 
Ch. dilepis and Ch. basiliscus, to be described immediately, one is 
disposed to believe that greater contractility in the case of the 
two latter may account for the great difference which they show 
from the species here under consideration ; especially since Milani’s 
figure * of the lung of Ch. basiliscus indicates long slender diverti- 
cula with slightly pronounced dilatations at their extremities. 

Chameleon basiliscus has lungs which have been described by 
Milani? and which agree most nearly perhaps with those of 
Ch. dilepis. The lung “itself is extensive and reaches back nearly 
to the kidney. The specimen which I had the opportunity of 
examining had been preserved for some time in alcohol. In 
neither lung could I find any ceca depending from the ventral 
margin of the organ, and in the left lung I did not find more 
than a single cecum at the posterior end of the lung, but conspi- 
cuous enough when detected by its yellow colour as contrasted 
with the colourless and transparent wall of the lung itself. In the 

* Zool. Jahrb. (Abth. f. Anat.) vil. p. 577. 
+ Loe. cit. p. 576. 


right lung, however, though there was a perfect agreement with 
the left lung i in the elhsemee of any ventral czecal outgrowths, such 
as occur in other Chameleons, there were three obvious ceca at 
posterior end. Milani figures five ceca. These were quite tubular 
and not swollen at the free extremity, as, for instance, in Ch. 
parvilobus. The walls are thick and they arise from a thick- 
walled portion of the lung. These ceca are, in fact, exactly like 
those of Ch. dilepis, from which the present species mainly differs 
in the extreme fewness of the ceca, as 1s apparent from Milani’s 

Chameleon pumilus has lungs which differ in several important 
points from those of the species that have been hitherto considered. 
In the first place, there are no signs whatever of any bronchi in 
the lungs. When the left lung is opened and the appearances 
presented compared with those to be seen in Ch. vulgaris, the 
following differences are recognisable. In both the aperture of 
communication between the two lungs, which represents, of course, 
the distal extremity of the bronchus, permits the interior of the 
right lung to be to some extent viewed. In the case of Ch. vul- 
garis the cartilaginous rings of the bronchus have to be cut up 
in order to display fully the aperture into the right lung through 
which are seen the cartilaginous rings of the bronchus of the right 
lung. In Ch. pumitlus, when the lateral wall of the left lung is 
removed no trace whatever of any bronchus is seen; there is 
simply a large circular orifice putting the two lungs into com- 
munication, which shows no traces of any bronchial cartilages that 
can be detected by the unaided eye. It is not plain whether this 
condition is to be regarded as primitive or as evidence of degene- 
ration. The lung itself is considerably shorter, relatively as well 
as actually, than in the species which has been dealt with in the 
preceding lines. It is, furthermore, different from the lungs of 
these other species in that the typical lung-structure persists 
throughout the whole sac. The alveoli in the lungs of Chame- 
leons generally are smaller and deeper proximally and get larger 
and shallower posteriorly, ultimately becoming pr: actically invisible. 
The hinder region of the lung is anangious. In Chameleon pumilus 
the alveoli become rather less marked posteriorly, but they are 
much more conspicuously circumscribed up to the very end of the 
lung than is the case with any of the larger species which I have 
had’ the opportunity of examining. The lung, in fact, is less 
metamorphosed into a mere air-sac in the present species than in 
any other which I have examined, excepting only Ch. tenio- 
bronchus, to which species I shall have to refer again immediately. 
In this particular it is plain that the lung of Ch. pumilus is more 
typically Lacertilian than that of such a species as Ch. vulgaris or 
Ch. calcarifer. 

A final peculiarity shown by the lung of this species is very 
remarkable. It has been stated in many general works that the 
Chameleons as a family are to be characterised by the cecal out- 
growths of the lungs, which have been considered in several 


species in the foregoing pages, and that is certainly the general 
impression among zoologists and anatomists. I was greatly sur- 
prised therefore to find that the lungs of Ch. pwmilus are quite 
unprovided with these otherwise characteristic outgrowths. The 
margin of the lungs is entire and slightly sinuous, the convexities 
occurring in the sinuous line being perhaps to be looked upon as 
rudiments or incipia of the cecal appendages. It will be observed 
that the absence of these ceca is associated with a more complete 
retention of the typical pulmonary structure of the lung, and 
therefore its greater efficiency as a breathing-organ. On the other 
hand, it is to be noted that where the cecal tubes exist the lung 
itself has lost considerably the alveolate structure and thus pre- 
sumably some of its efficiency as a breathing-organ. The Ophidia 
particularly show that the lung may be too large for its office as 
a respiratory organ, and they, like the Chameleons, are often 
lethargic in habit. 

The above account of the lungs of Chameleon pumilus is, 1 so 
far as the absence of tags is concerned, in harmony with the 
description of both Meckel* and Cuvier tf. The latter observes : 
“ Le Caméléon nain n’a rien de pareil; ses poumons sont deux 
petits sacs simples, ovales, de grandeur égale, comme ceux de la 
plupart des Sauriens”; and on another page: “ Ils manquent 
appendices.” Milani, however, obviously doubts these state- 
ments in writing £ as he does: “‘ Ob bei Chameleon pumilus die 
Ziptel wirklich fehlen, oder ob diese Behauptung nicht vielleicht 
auf ein mangelhaftes Praparat zuriickzufiihren ist, wage ich hier 
nicht zu entscheiden.” It is because of the latter doubt cast upon 
the facts that I have entered into the matter at some length, and, 
as I hope, settled it. 

I have finally to add to the description of the lungs in various 
Chameleons that Ch. teniobronchus agrees entirely with Ch. 
pumilus in the total absence of diverticula, an agreement which is 
very significant in view of other facts. 

The pigmentation of the interior of the body varies among the 
species of this genus. In all that I have examined the intestinal 
tract is a deep black, and there are generally (but not in Ch. ver- 
rucosus) patches of pigment upon the stomach not distributed so 
universally. The mesenteron is largely pigmented anteriorly 
in Ch. verrucosus. There is no variation, however, in the pig- 
mentation of the gut. The parietal walls are not so generally 
pigmented. It is, indeed, only in Ch. pumilus and in Ch. tenio- 
brenchus, among the species which I have examined, that the 
whole of the lining peritoneum of the body is of a deep black, 
quite as deep as is the gut. This pigmentation also extends to 
the mesenteries. In all of the remaining species the pigmentation 
of the general body-cavity and the mesenteries is hardly to be 
seen and only exists in very slight degree, so as not to affect the 

* “ Respirationsystem der Reptilien,’ Deutsch. Arch. f. d. Phys. 1818. 
+ Lecons @ Anat. Comp. 2me éd. par Duvernoy, t. vil. (Paris, 1840). 
ft Zool. Jahrb. (Abth. f. Anat.) vii. p. 573, footnote. 


general appearance. This peculiarity at once divides the two 
species mentioned from the rest, and other anatomical peculiarities 
described in the present communication tend to show the sepa- 
rateness of these two Chameleons from others. 

It may be remarked that the table of external characters used 
by Boulenger in the discrimination of the species of the genus 
brings together Ch. pumilus and Ch. teniobronchus*. 

Pancreas.—The shape of this organ shows differences in the 
species of Chameleon which I have examined. In all it lies 
partly between the stomach and the recurrent loop of the duo- 
denum, and partly dorsal of the stomach and to the posterior side 
of that organ. ‘That is to say, when the reptile is dissected and 
viewed in the ordinary position lying on the right side part of the 
pancreas, that lying between the stomach and the duodenum is 
visible and the rest is seen when the stomach is raised. The main 
differences in form are the relative thickness of the gland and the 
relations of the splenic lobe, which here, as in other Lizards, is to 
be distinguished at least to some extent from the rest of the gland. 
The distinction between the two lobes of the pancreas is most 
plainly to be observed in Ch. dilepis, where the splenic lobe is 
quite at right angles with the rest of the gland, and the duodenal 
part is continued on for a very short distance before it gives off 
the splenic lobe. In all the remaining species there is no such 
marked distinction, the two lobes forming one curved elongated 
mass. ‘This is particularly plain in Ch. teniobronchus, where the 
coils of the intestine lie entirely behind the pylorus, and the 
pancreas is therefore exposed for its whole length and not partially 
hidden by the stomach. I shall recur later to the coiling of the 
intestine in this and other species of Chameleon. 

The bulk of the gland differs greatly in the several species. 
In some it is much thinner than in others, and therefore, as the 
length is not far from being the same, relatively as to the size of 
the species the actual bulk fluctuates. Two extremes are well seen 
in the two species Ch, dilepis and Ch. calearifer, which, on account 
of their practically identical size, show the facts very plainly. In 
Ch. dilepis the gland is very thick, quite as thick as the diameter of 
the adjacent pyloric region of the stomach; its greatest diameter 
is about 6 mm. On the other hand, in Ch. calcarifer the pancreas 
is comparatively quite excessively slender, and only measures 
3 mm. or so in transverse diameter in the region which lies 
ventrally and in front of the stomach. There are similar differences 
between other species; but I do not give details, as the indi- 
vidual species vary so much in size that a comparison of the glands 
would involve rather complex measurements ; these would be of 
more value if the number of individuals examined were large. 
The prominent and easily recognisable differences between the 
two species selected will serve as an example of what also occurs 
elsewhere in the genus. There are, however, too great a series of 

* Cat. Lizards Brit. Mus. vol. 11. 1887, p. 440. 


gradations between the extremes to permit of the use in classi- 
fication of the dimensions of this gland. 

Liver-lobes—The proportions of the two lobes of the liver * 
differ markedly in several species of the Chameleons reported 
upon in the present communication. Thus in Ch. pumilus the 
right and left lobes are so nearly equal that only one can be seen 
with just traces of the other when the viscera are viewed from the 
left side. The gall-bladder is partly covered by the extensive left 
lobe, which, moreover, comes into contact with the stomach. 

The most extremely opposite conditions to these are shown (so 
far as the material in my hands enables me to say) in Oh. calearifer. 
In that Chameleon the left liver-lobe is very much shorter than 
the right. When the animal is viewed in the same posture as the 
last, the left lobe leaves exposed a section of the right lobe as long 
as itself, and does not even reach the gall-bladder, which lies on 
the right lobe not very far from its tip. There is, of course, no 
contact between the left lobe of the liver and the stomach, the 
ventrally flexed region of which lies considerably behind the end 
of even the right liver-lobe. 

In Ch, basiliscus the viscera in question are arranged and have 
very much the same proportions as in Ch. calearifer. The same 
may also be said of the Common Chameleon (Ch. vulgaris) and of 
Ch. verrucosus*. In Ch. parvilobus the two lobes of the liver are 
approximately equal, and the left lobe completely conceals the 
gall-bladder when that viscus is viewed from the left side. On 
the other hand, it, the left lobe, does not come so near to the 
stomach as in Ch. pumilus. Very much the same description will 
serve for Ch. dilepis, save for the fact that in this Chameleon the 
gall-bladder is not so completely hidden by the left lobe as in 
Ch. parvilobus and Ch. pumilus. The characters of the liver, 
therefore, hardly allow of any grouping of the species; for there 
are gradations. The alimentary canal does, however, show certain 
differences which permit of a grouping such as has been already 

In Ch. calcarifer and the other larger species the intestine is as 
well coiled as in other Lacertilia, and when the animal is opened 
from the side a good deal of the small intestine is seen to lie in a 
coil with secondary convolutions in front of, 7. e. headward of, the 
pyloric end of the stomach. The stomach, in fact, partly covers 
a section of the small intestine when the viscera are viewed from 
the left side. In Ch. pwmilus there is only one short bend of 
duodenum, which lies in front of the stomach. But the opposite 
extreme to Ch. calcarifer is to be seen in Ch. teniobronchus. In 
this small Chameleon the end of the stomach is not bent upon 
itself at all, but is continued back in a straight line to join the 
intestine, which is but little coiled upon itself. Moreover, the 
whole length of the intestine lies completely behind the stomach. 

* The liver itself is very compact in these reptiles, unlobulated, and with very 
firm outlines. 
+ I could see no gall-bladder in this species. 


There is thus a simplification in the coiling of the gut in this very 
small species which is not so strongly marked in the rather larger 
but still small Ch. pumilus. It is not without interest to note 
this apparent relation between smallness of size and simplification 
of structure shown also in the lungs of these species, as has been 
already commented upon *. 

(2) Some Notes upon Chameleolis. 

This Lizard is placed among the Iguanide in spite of its super- 
ficial likeness to a Chameleon. Indeed this superficial likeness is 
not after all very striking, and depends mainly upon the fact that 
the head is prolonged behind and above into a parietal crest. 
Nevertheless it 1s of advantage to be able to record a few facts in 
the visceral anatomy which distinctly confirm the placing of this 
genus in the immediate neighbourhood of Jguana. It has, more- 
over, its own peculiarities as compared with that genus; and 
therefore as a contribution to the visceral anatomy of the Lacer- 
tilia I am laying before the Society such facts as I have gathered 
from a dissection of a female individual. The existing knowledge 
of the anatomy of the Lacertilia shows that there are four 
marked structural features in which all the Iguanide that have 
been examined agree with each other, and the combination of 
which allows them to be defined. I shall, therefore, first of all 
deal with these four points, which together prove that Chame- 
leolis has been rightly placed among those Lizards. 

In the first place, the umbilical hgament which divides the two 
liver-lobes and is attached to the ventral median line of the body 
is a single ligament which runs continuously from end to end of 
the liver, without any trace of a posterior division upon the liver, 
as I have lately figured in Jguanat. The gall-bladder is left to 
the right of this hgament. In these particulars the umbilical 
ligament of Chameleolis is precisely like that of Jgwana, and there 
is no need to illustrate the relations of the ligament by a figure. 
I may mention that it is deeply pigmented. 

A second feature, which, though a small character, appears to be 
a constant one, is the position of the intercostal arteries in relation 
to the vertebre. In Chameleolis, as in some other Iguanoids, 
these arteries plunge into the thickness of the dorsal parietes 
towards the posterior end of each vertebra; in some other Lizards 
they disappear from view at about the middle of each vertebra. 
Of course, Chamcleolis has the same regular arrangement of pairs 
of these arteries as in other Lizards, a feature, indeed, which seems 
to differentiate the Lacertilia from the Snakes, at least broadly 

Thirdly, Milani £, whose account of the Lacertilian lungs is the 
most recent and comprehensive known to me, has found that in 
the Iguanide the lung is totally divided into two chambers, of 

* Supra, pp. 39 & 41. 

+ P.Z.S. 1905, vol. i. p. 12, fig. 7. 
{ Zool. Jahrb. (Abth. f. Anat.) vii. p. 545. 


which the more dorsal extends headwards of the orifice of the 
bronchus. This statement at any rate holds good for the majority 
of the Iguanide that have been examined. Chameleolis does not 
agree with these types, for the lung is not divided by an obliquely 
placed septum into two approximately parallel chambers. It is, 
nevertheless, not to be removed from the Iguanide on this 
account, since it appears to present points of likeness to the 
undoubtedly Squamoid Phrynosoma, and one point at least of 
resemblance to the Iguanoid genus Polychrus. 

Text-fig. 13. 

Lung of Chameleolis, opened longitudinally. 

The bronchus enters for a short distance into the lung as a 
completely circular tube; there is no snake-like series of flattened 
semirings such as is to be found in Jguana*. The projecting 
bronchus is, as in Phrynosoma, moored to the walls of the lung by 
septa. The cavity of the lung, therefore, extends headwards of 
the opening of the bronchus and all round it. There is no septum 
in either lung which separates off the dorsally placed caecum of the 
lung as a distinct cavity from the rest of the cavity of the organ. 
In the left lung the structure happened to be more favourable for 
observation than the right lung, and I have accordingly had a 
drawing prepared (text-fig. 13) of the interior of this lung. It 

* Milani, Joc. cit. pl. xxxi. fig. 13. 


will be there seen that the strong septa which produce a pouching 
of the dorsal region of the lung in other Iguanids and Agamids 
are also to be seen in Chameleolis. Lobserved six of the chambers 
altogether, of which three would appear to belong to the anterior 
part of the lung, 7. e., that region which is in other Iguanids 
divided off by a septum from the posterior region, and three larger 
pouches belonging to the posterior region of the lung. Finally, 
the end of the lung abruptly narrows and forms a finger-shaped 
region with a but slightly marked network. It seems to me to be 
possible to compare this with the Chameleon-like outgrowths of 
the lung in Polychrus marmoratus *. 

In the fourth place, the right extremity of the liver is attached 
by a fold of membrane which separates the lung from the post- 
hepatic region of the body-cavity and is continuous with the 
oviducal membrane. 

Besides these points, which, together with various external and 
osteological characters used by others, fix the systematic position 
of Chameleolis, there are other features in its anatomy which I 
have ascertained and which are worth noting as a contribution to 
Lacertilian structure. 

The pigmentation of the body-cavity is in some ways remark- 
able. The umbilical ligament, not only the region which is 
attached to the liver, but that which is attached to the stomach, 
is deep black, and in the latter region contrasts with the yellowish 
gut. The gut itself is, however, pigmented in the case of the 
large intestine. This pigmentation is limited to the dorsal side 
of the gut and involves the whole of the cecum, The appearance 
presented is, indeed, of two tubes closely applied, of which one is 
the small intestine and the other ends at the blind extremity of 
the cecum. _ 

As in many Lacertilia, the peritoneum generally is deeply pig- 
mented, and a distinction is to be drawn between the posterior 
pigmented region and the anterior region of the body-cavity, 
where its walls are not pigmented at all, so far as naked-eye 
appearances go. 

While, however, in most Lacertilia this line of demarcation is 
quite oblique, bending ventrally in a continuous curve, it is in 
Chameleolis quite transverse (to the longitudinal axis of the body) 
in direction, but with a curved outline, now convex, now concave. 

The existence of bundles of plein muscular fibres in the mes- 
enteries reaches a very great degree of development in many 
Lacertilia. In the Lizard which forms the subject of the present 
communication there was no development of such fibres that could 
be seen with the unaided eye. The ovaries contained no mature 
ova. There was a fully formed egg in each oviduct, with a dirty 
white shell of leathery consistency. There was no trace of an 
embryo in the egg. 

The apex of the heart is fixed to the pericardium by a very 

* Milani, loc. cit. pl. xxxi. fig. 15, 


slender gubernaculum cordis, a structure which is rarely absent 
from the heart of the Lacertilia *. 

The arterial system presents certain peculiarities as compared 
with that of other Lacertilia. The disposition of the intercostal 

Text-fig. 14. 

B ~~ -Coel. 

A. Stomach and pancreas of Chameleolis. 
P. Pancreas; p.v. Portal vein; p.v.g. Gastric portals; Sp. Spleen. 
B. Dorsal aorta and branches of the same. 

Cel. Coeliac artery ; i. Intestinal arteries; ws. @sophageal arteries ; Sp. Spleen. 

arteries has alveady been mentioned as a point of affinity with the 
Iguanide. The visceral arteries (text-fig. 14) which supply the 

* But is absent m Varanus occasionally (see Beddard, P. Z.S. 1906, vol. ii. p. 617 
footnote) and Zonurus giganteus (infra, p. 55). 


alimentary canal are collected into two groups, leaving a tract of 
considerable length from which no arteries to the gut arise. The 
anterior group is situated just behind the union of the two aortic 
arches, and consists of no less than seven small arteries supplying 
the cesophagus and stomach, ‘These arise from both sides of the 
aorta and are partly arranged in pairs; they run to both sides of 
the stomach, There is then a long gap until the origin of the 
intestinal arteries. The general ‘plan of these is like that in 
most Lacertilia ; but there are differences in detail from those of 
many genera, When the mesentery is turned over to the right 
the cecal arte xy which arises most anteriorly is seen to run over 
the following duodenal artery, but under the third artery, that 
which : supplies the spleen, &e, 

With regard to the venous system, the only notes that I have 
made refer to the hepatic por tal system. The junction of the 
anterior abdominal and the main portal trunk is very near to the 
conjoint entrance of both into the liver—much nearer than is the 
case with many Lizards. In addition to this, the chief portal 
trunk, there are two vessels which pour blood direct from the 
stomach into the liver (text-fig. 14): one of these, the more 
posterior in position and the larger, is associated with the left 
lobe; the other, a slender twig, enters the liver to the right of the 
last described. The ventral parietal hepatics are also two, of which 
one is a little to the right of the other in its point of entrance to 
the liver. Both are rather far forward on the liver. There is 
but one dorsal parieto-hepatic. This, as is the case with other 
Lizards*, is associated with the “ Hohlvenenfortsatz” of the liver, 
and runs in the mesentery, binding that lobe of the liver to 
the right parietes. It runs a conside ‘able way forwards along 
the ver rtebral column before becoming lost in the thickness of the 

The pancreas of Chameleolis (text-fig. 14) i is constructed upon 
the usual Lacertilian plan, but differs in various details from that 
of other Lizards. It isa Y-shapel gland and completely solid 
throughout. There are no thin diffuse branches spread through 
the mesentery such as are to be found in the case of the pancreas 
of Zonurus giganteust. One arm of the ¥ ends, after dilating 
slightly, in the concavity of the somewhat bean- shaped spleen ; the 
other forms a thick mass in contact with the commencement of 
the duodenum. The stem of the Y forms a thin rod of pancreatic 
tissue, which closely accompanies the portal vein and very nearly 
touches the liver. This region of the pancreas seems to me to be 
longer than in some other Lacertilia, though in most there is a 
process of the pancreas running in the same direction. The 
splenic lobe of the pancreas is not extraordinarily thin, as it is in 
Tiliqua scincoides ~, but of fairly robust diameter, 

* The absence of this vessel is rare, but Hochstetter, whom I have been able to 
confirm, has asserted its absence in Chameleon vulgaris. I take this opportunity 
of stating that this vein is also absent 1 in Chameleon verrucosus. 

+ Vide infra, p. 55. { See Beddard, P. Z. §. 1905, vol. ii. p. 262 

Proc. Zoou. Soc.—1907, No. IV. 4 


(3) The Position of the Umbilicus in certain Vipers. 

IT am not aware that the point of entrance of the umbilical sac 
into the body in Snakes has ever been made use of as a systematic 
character. I find, however, from a few observations that I have 
been able to make recently, that this anatomical relationship is 
apparently of systematic value. Since of one species, viz. Lachesis 
lanceolatus, selected for these observations, I have been able to 
examine a considerable number of individuals, the variation of 
the character from one individual to another became a matter of 
additional interest, especially in view of the fact that all the indi- 
viduals were of one brood. It appears that in Vipers, as compared 
at any rate with the Anaconda*, the umbilicus is much nearer tothe 
cloacal aperture. I have examined fourteen individuals of Lachesis 
lanceolatus of the same brood and of approximately the same size, 
though they died on different dates, from March the 9th to May. 
The length varied from 113 to 12 inches exclusive of the short 
tail. I do not give measurements in millimetres, since to use such 
gives an appearance of rigid accuracy not attainable in a dead 
snake capable of artificial extension and shortening. In nearly 
all of these fourteen individuals four scales occupied the umbilical 
region, each of them being bisected by a groove running longi- 
tudinally to the axis of the body. I found, in fact, that there 
were four scales thus modified in eleven individuals. In two of 
the remaining snakes there were five of these scales in which the 
two sides had not joined across the middle ventral line, and in 
the fourteenth individual only three scales and a portion of the 
fourth ; the number of scales intervening between the last of the 
‘‘ umbilical” scales and the anal scale varied a good deal but within 
very narrow limits. The actual facts are these: in three speci- 
mens 17 scales intervened between the points mentioned ; in one 
specimen 18 scales; in five others 19 scales; in three 20 scales; 
in one 21 scales; and, finally, in one 22 scales. The average is 
thus arithmetically 19, and actually there were more specimens 
exhibiting the average than any other number. Having due 
regard to the narrow.range of the variation, it seems likely that 
the position of the umbilicus in this species of Viper can be 
regarded with safety as lying 19 scales in front of the anal scale. 
It is important to notice the length of time during which this 
feetal character is retained. The last specimens examined by 
myself died on May 15th of last year. These and the other 
individuals were acquired by the Society on Dec. 12th, 1905. 
The last specimens examined by me were therefore more than six 
months old. J have some confidence, therefore, in comparing 
Lachesis lanceolatus in respect of these characters with other 
Vipers of an obviously greater age. I may first of all, however, 
refer to newly-born Vipers which I have recently dealt with tf in 

* See Beddard, P. Z.S. 1906, vol. i. p. 13. 
+ P.Z.S. 1906, vol. i. p. 34. 


a paper communicated to this Society. It is impossible to be 
certain of the exact position of the actual umbilicus in Lachesis 
lanceolatus for the purposes of comparison between these two 
types, 7. e., which of the four or five broken scales correspond to 
the two scales in Bitis nasicornis which are actually divided by 
the foetal blood-vessels. Assuming, however, that they are even 
the last two, there still remains a substantial difference in position 
between the umbilicus of the twospecies. For in Ditis nasicornis 
the actual numbers of scales intervening between the umbilicus 
and the anal scale are respectively in the five examples studied 9, 
11, 11,12,14. There is thus exactly the same amount of variation 
as in Lachesis lanceolatus, but round a different mean. 

The position of the umbilicus in Russell’s Viper (Vipera 
russellit) is again different and relatively more fixed than in either 
of the species hitherto considered. As in Sitis nasicornis, the 
actual umbilicus consists of two scales only, which do not meet 
ventrally, and between which the plug of tissue bearing the 
umbilical vein &e. passes into the interior of the body. The 
young Vipers in question were a very few days old, but all 
external traces of the yolk-sac had disappeared. In front of 
these two scales either two or three scales were divided by a suture 
in the middle ventral line, and posteriorly to the two ‘“ umbilical 
scales ” either one or two scales were similarly split by a ventral 
suture. Between the last of the two completely divided scales 
and the anal scale there intervene in the five examples examined 
respectively 16, 16, 16, 17, 17 scales. ‘The position of the umbi- 
licus is therefore different in this Viper, and its fluctuations of 
position are less than in the two species to which I have already 
called attention. It is perhaps permissible to call attention to 
the fact that Vipera and Lachesis agree with one another more 
nearly than either does with Sitis. This is, of course, not in 
accord with generally received views upon the classification of 

J have examined several Vipers of more mature age, and in 
two specimens, at any rate, I find what appear to be obvious traces 
of the umbilicus. In a not fully-grown example of itis arietans 
measuring 30 inches from the snout to the cloaca, four scales 
showed a line of division in the ventral median line. The second 
of these had the most strongly marked groove, and_ possibly 
therefore represents one of the two scales already described in 
the young as immediately surrounding the stalk of the yolk-sac. 
Between the last of the grooved scales and the anal scale 9 scales 
intervened. ‘The species evidently therefore comes nearest to the 
other species of Sitis which has been described above. Inasecond 
specimen measuring 32 inches there were 12 scales between the 
last of four grooved scales and the anal scale. I have also seen 
similar traces in a large adult example of Bitis gabonica. Here 
there were also four scales showing traces of the umbilicus; but 
instead of being grooved they were merely nicked posteriorly. 
Between the last of them and the anal scale 8 scales interyened 



Tt is not safe upon these two last-mentioned examples to attempt 
to draw any distinctions between the different species of Bitis. 
It seems, however, to be most probable that they do not differ 
widely from each other as regards the points under discussion, 
whether they will ultimately be found to differ specifically or not. 
It is, however, quite plain, in reviewing all the facts brought 
forward in the present communication, that the position of the 
umbilicus among Vipers is one that does at least characterise 
some forms which happen in the instances studied to be generi- 
cally separated. 

(4) Some Notes upon the Anatomy of Zonurus, with Special 
Reference to the Hyoid. 

The following notes refer to three specimens of Zonurus 
giganteus which I have had the opportunity of dissecting during 
the last year or two. The anatomy of this Lizard is already to 
some extent known through the work of previous observers. 
The lungs have been dealt with by Milani* in his general account 
of these organs among the Lacertilia, and the arteries of the 
gastric and intestinal regions are described and figured by Hoch- 
stetter +. There remain, however, a few points to which it is 
worth while calling attention as a further contribution to the 
natural history of this Lacertilian. 

Of special importance—rather, however, from a general point 
of view than as a particular contribution to our knowledge of this 
Lizard—is the condition of the elements which together make up 
the hyoid complex of bones and cartilages in this Lacertilian. 
T am able to add to what I have to say concerning Zonurus a few 
notes upon other genera of Lacertilia which I have dissected for 
purposes of comparison. I commence with a brief résumé of 
some of the facts already known of this part of the skeleton. 

The hyoid and branchial arches of Lacertilia have not, as it 
appears, been investigated in a very large number of genera. 
Several are figured in the volumet of Bronn’s ‘'Thierreich’ dealing 
with the Lacertilia, while other genera have been illustrated by 
subsequent writers §. Apart from differences in the form of the 
individual elements of the hyoid complex there is substantial 
agreement, according to these various writers. For contrary to 
what is to be found in the Chelonia—where the remains of the 
hyoid arch proper is followed by two branchial bars considerably 
developed—the Lacertilia are generally believed to be characterised 
by the preservation in the adult of only one visceral arch follow- 
ing the hyoid arch, which is stated to be the first branchial. 
This statement occurs at any rate in such authoritative textbooks 

* Zool. Jahrb. (Abth. f. Anat.) vil. p. 545. 

+ Morph. Jahrb. xxvi. 

+ Reptilia, Bd. vi. Abth. iii. Taf. 72. figs. 2-8, & Taf. 107. figs. 24, 33. 

§ E. g., Gecko mauritanicus, Gadow, Phil. Trans. 1888 B, pl. 72. fig. 10; Helo- 
derma suspectum, Shufeldt, P.Z.S. 1890, pl. xviii. fig. 6; Chlamydosaurus and 
Physignathus, Beddard, P. Z. 8. 1908, vol. 1. p. 20, text-fig. 9,and p. 21, text-fig. 10. 


as those of Hatchett Jackson *, Gadow 7, and Sedgwick ¢, which 
may be regarded as expressing the current knowledge of the 
subject. Nevertheless, the late Prof. W. K. Parker$, in describing 
the adult skull of Lacerta agilis, wrote as follows :—‘‘ Another bar, 
half as long as the first, and unossified, lies behind the first 
branchial above; it is f-shaped, with the top hooked inwards, 
like the lower piece; this is the upper (0r.?), or ‘epibranchial’ 
part ; it has a small snag outside its middle. Besides this, there 
is, on each side, a slender, slightly outbent hypo-branchial (/.67r.); 
this belongs to the second branchial, and also from its length is 
evidently part of the third, neither of which chondrify, above, in 
the embryo.” In a footnote is added the remark that “this little 
highly-metamorphosed Lizard has scarcely thrown aside the 
skeleton of these organs of aquatic respiration.” It is obvious 
that Prof. Parker’s account is a little misleading, and this doubt- 
less accounts for the fact that the existence of remains of a second 
branchial arch in Lacerta has been largely ignored in zoological 
literature. What he speaks of as an “ epibranchial,” without 
determining to which arch it belongs, but letters “ br.*,” is clearly 
from its position a vestige of the second branchial arch, as is plainly 
recognised in Prof. T. J. Parker’s ‘Zootomy’ and in his ‘Textbook 
of Zoology,’ written in conjunction with Prof. Haswell|. The 
exceptional character of the hyoid complex of Lacerta in possess- 
ing “the epibranchial of the second branchial arch” is properly 
emphasised by Dr. Shufeldt {| in reviewing existing knowledge of 
the Lacertilian hyoid bones. 

The third postmandibular arch is, however, by no means a 
peculiarity restricted to Laverta. It occurs in Zonurus in the 
form of a short and slender bar lying behind the well-developed 
first branchial bar **. This bar of cartilage does not extend down 
to the median copula, and indeed falls a considerable distance 
short of it. I have examined three other Lacertilians in which 
this same visceral arch is represented and one in which it is not 
to be found; but I have not at present attempted an exhaustive 
research into the facts of its absence or presence among the 
different families of Lacertilia. I could not detect the bar of 
cartilage in Chameleolis, whose anatomy has been described above. 
Tt is well developed in both 7iliqua and Trachydosaurus. Inthe 
former (text-fig. 15, p. 54) it is very conspicuous, and it is nota 
little surprising that it has been missed, unless I have unwittingly 
overlooked its description somewhere. But if this be the case it is 
clear that its existence has escaped the writers of many textbooks. 

* Ond ed. of Rolleston’s ‘ Forms of Animal Life.’ 
+ “ Amphibia and Reptilia,” in ‘Cambridge Natural History.’ 
t ‘Textbook of Zoology,’ vol. 1. 
§ “Development of Skull in Lacertilia,’ Phil. Trans. 1879, p. 616. 
|| ‘Textbook of Zoology,’ vol. 11. 
q P.Z.S. 1890, p. 225. 
*k There is no trace of this shown in a figure of the hyoid of Zonurus cordylus 
copied from Henle in Bronn’s ‘ Thierreich,’ Reptilien, vol. vi. Abth. 11. Taf. 107. 
fig. 33. 


In Tiliqua this second branchial arch is more extensive, as it 
appears, than in Zonurus, and lies obliquely across the first 
branchial arch, though beneath it. The latter arch ends in a 
curled piece of cartilage which is directed backwards and overlaps 
the third post-mandibular arch. But the position varies according 
to the degree of distortion of the muscles of the neck of the reptile. 

Text-fig. 15. 

H Br. 

Tiliqua scincoides, head and neck. 
Dissected to show three postmandibular visceral arches in sitw. 
To the left the isolated extremities of the same arches in another individual. 

Br.1. First branchial arch; Br.2. Second branchial arch ; H. Hyoid arch. 

Towards the lower end of the bar is a triangular, projecting, 
“snag” (not visible in one of two examples dissected) like that which 
Prof. W. K. Parker has figured in Lacerta. To this projecting 
process is fixed a ligament which is inserted on the free dorsal end 


of the hyoid arch. The ligament thus avoids the first branchial 
under which it lies. It is not surprising to find that 7rachydo- 
saurus, so closely allied to Z'iliqua, also possesses this second 
branchial arch. As in the genera mentioned, the arch is only 
represented by its upper part, the epibranchial, as Parker termed 
the equivalent cartilage in Lacerta. Finally, I have to record 
that the bar of cartilage is also found in Gerrhonotus, a genus 
of whose anatomy I offer some further notes below. In the 
present state of our knowledge it is not possible to state whether 
or not this occurrence does or does not bear wpon the affinities of 
Gerrhonotus. The cartilage was not so easy to find in this small 
Lizard, where it is siender and delicate, but can be detected 
by gently moving in various directions the muscles in its vicinity ; 
the stiff ends of the cartilage thus become apparent. 

A second feature in the anatomy of Zonwrus to which I desire 
to draw attention is the total absence of the gubernaculum fixing 
the apex of the ventricle to the walls of the pericardium. This 
ligamentous band or thread (it varies in importance in different 
genera) is so usual among the Lacertilia as to be characteristic of 
that order of Reptiles, as it is, indeed, of others. I have already 
pointed out that the gubernaculum cordis is not to be found in 
the heart of Varanus niloticus and some other species*. It is 
interesting to notice that this absence of the gubernaculum which 
is universal in the higher Vertebrates (Aves and Mammalia), as 
well as generally in the Ophidia *, is sporadically developed among 
the Lacertilia. It should also be mentioned that this condition 
of the heart was found in two examples of Zonurus giganteus (the 
third was not examined ad hoc)t; it is therefore probably charac- 
teristic of the species if not of the genus. 

The liver of this Lizard is unusual in its form. The right lobe 

is prolonged in the usual way over the vena cava. But the left 
lobe, instead of being but slightly divided at the entry of the 
anterior abdominal vein, is deeply bifid thereat §. The whole 
organ is thus markedly trifid posteriorly and is not unsuggestive, 
in appearance, of the mammalian liver. 

The pancreas displays one noteworthy character. Its general 
form is like that of the majority of Lacertilia. The organ embraces 
the stomach, being found on both sides of it; the splenic lobe is 
fairly stout and reaches the spleen, and there is a process of the 
gland extending towards the liver. The peculiarity of the pancreas 
of this Lizard is that diffuse thin ramifying tags of pancreatic 
tissue lie in the mesentery on either side of the splenic lobe of the 
gland with which they are connected. This tendency towards a 
diffuse irregularly shaped thin pancreas is obviously to be com- 

pared with the conditions obtaining in the Chelonia. 
* P, Z.S. 1906, vol. ii. p. 617 footnote. 
+ Perhaps universally also. In any case the occasional ligament tying the apex 
of the heart to the pericardium is rather different (see P. Z.S. 1904, vol. ii. p. 107). 
+ I have since found the same absence of the ligament in another example. 

§ I am not quite certain that it is not the right lobe which is thus bifid. It is a 
point difficult to settle. 


(5) Some new Facts bearing upon the Affinities of 

This genus of Lacertilia is sometimes placed in a special family 
of Lacertilia, the Gerrhonotide. By others (¢. g., Boulenger *, 
Gadow *) it is relegated to the Anguide. The general aspect otf 
Gerrhonotus ceruleus is, on the other hand, by no means unlike 
that of the Scincide ; I am not aware that any notes have ever 
been published upon the visceral anatomy of this Lizard. I 
venture, therefore, to lay before the Society some notes which a 
recent dissection of more than one example of Gerrhonotus ceruleus 
enables me to offer as a contribution towards the determination 
of the systematic position of this genus or representative of a 

T have by no means attempted an exhaustive survey of the 
anatomy of this Lizard. But J] am able to note down a few facts, 
all of which are of some interest from the point of view of a 
comparison with other Lacertilia. The structure of the guadrato- 
jugal ligament is one of the characters which I carefully examined 
in Gerrhonotus. I find that the arrangement and appearance of 
this ligament is precisely as it is in the genus Gerrhosaurus and 
in the Skink Humeces~. That is to say, the ligament of this 
genus is very distinctly marked off and of equal breadth through- 
out, nowhere vaguely shading off into surrounding tissues. 
Moreover, it is attached on the one hand, of course, to the quadrate 
bone and on the other to the bony scales which cover the face in 
this region. It is not inserted on to any bone of the skull. In 
the present state of our knowledge it is not possible to comment 
upon this likeness to Gerrhosaurus and Hwmeces as an argument 
in favour of the Skinkoid affinities of Gerrhonotus, though I have 
thought it worth while to record the fact for future comparison. 
The second feature in the structure to which I draw attention is the 
complete pigmentation of the interior of the body. There is here 
no paler area divided by the oviducal mesentery from a more 
darkly pigmented posterior portion. 

As is now well known §, the umbilical hgament of the Skinks 
is frequently a double ligament attached to the ventral surface 
of the liver along two parallel lines which become confluent 
anteriorly. I observed nothing of the kind in Gerrhonotus, where 
the umbilical ligament is, as in most Lizards, a single mesentery. 
Tn this anatomical fact there is a likeness to Ophiosaurus as well 
as, of course, to Zgwanaand other Lizards. In any case the Lizard 
shows no affinities to the Scincide. 

The pancreas and the spleen and their relationship to one 
another differ greatly among the Lacertilia, and more than one 
fact in the structure of the two glands is recorded in the present 

* Catalogue of Lizards in the Collection of the British Museum. 
+ “Reptilia,” in ‘Cambridge Natural History,’ p. 538. 

t P. Z.S. 1905, vol. ii. p. 256. 

§ Beddard, P. Z.S. 1888, p. 102. 


communication *, In Gerrhonotus the splenic prolongation of the 
pancreas is present, but it does not reach the spleen at all, though 
extending a good way in the direction of that organ. Among 
the Skinks this pancreatic process towards the spleen is to be 
found, as I have already recorded‘, in the genus J%liqua, and can 
confirm in all details from a subsequently examined example of 
that genus. 

There is, however, no particular likeness in the structure of the 
pancreas of Gerrhonotus to that of Ophisaurust. In the latter 
the pancreas consists only of two closely applied lobes which rest 
upon the ventral surface of the pylorus and small intestine, there 
is no vestige of a splenic lobe§. The spleen of Gerrhonotus is 
rather peculiar in position. Very generally among the Lacertilia 
this blood-gland is elongated and somewhat bean-shaped in out- 
line, and hes with its long axis parallel with the long axis of the 
stomach. In Gerrhonotus the shape is quite normal, but the long 
axis 18 perpendicular to the long axis of the stomach. 

The hepatic portal system of veins of Gerrhonotus ceruleus 
varied but little in the two specimens dissected. The ventral 
parieto-hepatic veins running in the umbilical ligament were 
three in each Lizard. The first two crossed each in their course 
in one specimen, and perhaps in both, though I have no note as 
to this in the second example dissected. ‘The crossing is such 
that the anterior of the two vessels draws blood from a region of 
the ventral body behind that which is supplied by the posterior 
of the two veins. 

The dorsal parieto-hepatics are either two or three and are 
otherwise quite normal in their position. The arrangement of 
the gastro-hepatic veins is interesting in relation to the question 
of the affinities of the genus Gerrhonotus. There are either four 
or five of these vessels of somewhat varying calibre arranged close 
together, and thus forming a ladder-like structure lying quite at 
the anterior end of the liver and running to this from the adjacent 
border of the stomach. There are no gastro-hepatic veins situated 
more posteriorly. The interest attaching to this arrangement of 
the vessels is that it is completely paralleled in Ophisawrus apus ||, 
making allowances for the greater elongation of the liver in the 
latter snake-like Lizard. 

In Ophisaurus, in fact, there are six of these veins. Now, as a 
rule, the Lacertilia have not a great many separate gastro-hepatic 
vessels. J have myself examined several species embracing as 
many genera and find the following facts, some of which I have 

* Cf. pp. 48 & 55. + P.Z.S. 1905, vol. ii. p. 262. 

{ In contrast to this difference in form between the pancreas of genera which 
appear to be allied is the close resemblance in another case which I take this oppor- 
tunity of recording. In both Iguana tuberculata and Liolemus magellanicus (I 
owe the specimen to the kindness of my friend Capt. Richard Crawshay), which are 
both Iguanide, but not much alike superficially, the long splenic lobe of the pancreas 
just touches the posterior end of the spleen. 

§ See P. Z.S. 1905, vol. ii. p. 4.75, text-fig. 64. Anguis also lacks the splenic 

|| See P. Z. S. 1905, vol. ii. p. 475, text-fig. 64. 


already made known in recent communications to the ponely 
upon the anatomy of these Reptiles. In Jgwana tuberculata I 
found in two examples two gastro-hepatic veins, and precisely the 
same arrangement characterised two examples of Amphibolurus 
barbatus. Uromastix acanthinurus showed, also in two examples, 
a single vein, which, however, was made up of three considerable 
affluents from the stomach ; these, it will be understood, entered 
the liver as a single vessel. In one of the specimens the third 
affluent only joined the common trunk formed by the other two 
just before their entrance into the liver. In Gerrhonotus, Tupi- 
nambis, Chameleon, Phelsuma, Tarentola, | have recorded, or am 
now able to record, the existence of only one gastro-hepatic vein, 
which however is, as a rule, made up of two affluents. The 
Scincide form an exception to the general arrangement of these 
vessels, and at first sight appear therefore to be near akin in this 
particular to Gerrhonotus. 

Of Tiliqua scincoides | have dissected two examples for the 
purposes of the present investigation, and find in both of them 
the following arrangement of the gastro-hepatic veins. There 
are four of these, which appear at first sight to lie accurately side 
by side in the gastro-hepatic mesentery. A more careful exami- 
nation, however, shows that the stomach is bound to the liver by 
two mesenteries, one above the other, as seen when the animal is 
opened along the median ventral line and the viscera examined 
in an tndissmbed condition. The lower of these, i.e. that which 
lies above in the ordinary position adopted in dissection, is the 
gastro-hepatic mesentery found in all Lizards. When this is cut 
through a second mesentery comes into view, which is attached to 
the right side of the liver and to the more dorsal side of the 
stomach. This mesentery exists in other Saurians, but is inserted 
on to the mesogastrium and does not touch the stomach at all. 
Whether this arrangement of the right dorsal suspensory hgament 
of the liver has anything to do with the double umbilical hgament 
of the same family of Lizards is not certain; but it is found in 
most but not in all Skinks. I find it in the genus which we are 
now considering, in Seps (Chalcides), Scineus, Hwmeces, and Macro- 
scincus. It is not to be found in Trachydosaurus rugosus. To 
revert to the gastro-hepatic veims in Z'%liqua scincoides, the most 
posterior of the four veins runs along the right or lower (as seen 
on dissection) gastro-hepatic mesentery; in front of it are two 
of the veins which run in the upper or left gastro-hepatic mesen- 
tery (the mesentery present in all Lizards). Finally, there is a 
single vein which is inserted just at the junction anteriorly of the 
lines of attachment of the two mesenter ies, to the lower (dorsal) 
surface of the liver. In Trach ydasourus, which, although a 
member of the family Scincide, agrees with other Lizards in the 
presence of only one gastro-hepatic mesentery, I find in an 
example dissected only » one gastro-hepatic vein, which, as is so 
usual, is formed by two equally sized affluents, I have some 
notes, however, of an example, dissected a good many years ago, 



in which in addition to this there was another vein further 
forward at the junction of the two dorsal suspensory mesenteries 
of the liver. dMJacroscincus is normal—for a Skink—ain the presence 
of two gastro-hepatic mesenteries ; and yet it has only one gastro- 
hepatic vein. This is formed of two equisized affluents, and runs 
in the right-hand mesentery, the other being quite anangious. 

Eumeces algeriensis shows the same double series of gastro- 
hepatic veins that are to be found in Z%liqua scincoides. ‘There 
is one vein only in each of the two gastro-hepatic hgaments and 
a third vein implanted at the junction of these anteriorly. As 
in Tiliqua, the medianly situate vein of the three belongs to the 
left-hand ligament. Of an example of Chalcides ocellatus, dissected 
by me a good many years ago, I have sketches ae descriptions 
showing that this species is more like Macroscincus* than Humeces 
or Piligua. For the gastro-hepatic veins are limited to the right- 
hand one of the two gastro-hepatic ligaments, with the usual 
vein which enters the liver at the junction of the two veins. 
There were in this individual four of these veins in the right 
ligament. It seems, therefore, that the numerous gastvo-hepatic 
veins of the genus Gerrhonotus may be regarded as evidence of 
affinity with the Anguide, since these veins are, as a rule, not 
numerous among the Lacertilia other than the Anguide, Amphis- 
beenidee, and arent “ia, except among the cemenees where the 
existence of two castro- hepatic ligaments accounts for the greater 
number of these veins than occurs in the majority of those genera 
with only one gastro-hepatic ligament. And, in coming to this 
conclusion, it must be further borne in mind that the number of 
gastro-hepatic veins would appear to be fairly constant, so far as 
the somewhat meagre facts already known allow us to judge. 
There is but little alse in the anatomy of this Lizard, so far as I 
have been able to record the facts, which bears very distinctly 
upon its affinities, a conclusion for which the, at present, very 
small knowledge of the Lacertilia is doubtless largely responsible. 
It seems, at any rate, to be the fact that Gerrhonotus exhibits no 
marked features in its organisation which poimt to an affinity 
with the Scincide, except perhaps the condition of the quadrato- 
jugal ligament, which is undoubtedly like that of Huwmeces. 

(6) On a Point of Structural Resemblance between Heloderma 
and Varanus, and on some Specific Characters of Varvanus. 

Although it is not the prevalent opinion that these two genera 
of Saurians are nearly allied, there are not wanting anatomical 
resemblances between them; and, indeed, some of the most recent 
writers 7 on the anatomy of Heloderma have brought together a 

* It is perhaps not without interest to notice that in these two genera (Chalcides 
and Macroscineus) the double character of the umbilical ligament is not so marked 
as it is in Humeces and Tiliqua. They have, therefore, at any rate, two anatomical 

features in common. 
+ Boulenger, P.Z.S. 1891, p. 109; Beddard, ibid. 1906, vol. ii. p. 601. 



good many facts in favour of such an alliance, On the other 
hand, Dr. Shufeldt *, who has given us a comprehensive sketch of 
the anatomy of Heloderma, remarked that his own studies of the 
Varanide convinced him “of the fact that Meloderma is far 
removed from that group, having very little structural affinity 
with it.” To these papers cited below, and to others quoted in 
them, reference may be made for the views which have been 
held with regard to the position occupied by the genus Heloderma 
in the system. 

In recently dissecting examples of these two genera I have 
noticed two structural features in which the two genera are similar 
and by which they may be differentiated from any other Lacer- 
tilians whose anatomy is known, so far as concerns the points in 
question. The first of these is a feature in the anatomy of the 
respiratory organs which has indeed been described in Varanus 
but not in Heloderma. As to the former genus, Meckel 7, 
Giinther f, and Milani §, to whose investigations our knowledge 
of the anatomy of the respiratory organs in the Varanide is 
chiefly due, describe a short branch given off by the bronchus 
shortly after it has entered the lung; this supplies the headward 
extension of the lung which is so well marked in this genus of 
Lizards. It is plainly figured by Milani||, whose illustrations 4, 
particularly a diagrammatic figure, show that this twig arises im 
front of an aperture in the walls of the intrapulmonary bronchus. 
Some of Milani’s figures** also illustrate another somewhat im- 
portant fact, which is that the bronchus until it gives off the branch 
to which reference has been made does not really lie within the 
the lung, but outside of it. The lung in growing forward has 
wrapped round the end of the bronchus. Though apparently 
within, this portion of the bronchus is really morphologically 
outside of the lung. The interest attaching to the exact relation- 
ship between the bronchus, the lung-tissue, and the first branch 
of the bronchus appears to me to be this :—that this independent 
branch arising so early from the bronchus 1s possibly to be compared 
with the eparterial bronchus of the Mammalia. This comparison 
is not suggested by Milani. In any case it is, so far as I am 
aware, a structure that has not yet been described in any other 
Lacertilian. A precisely comparable branch of the bronchus 
occurs, however, in Heloderma, where its existence is an interesting 
feature of resemblance to Varanus. I cannot find that any of 
the writers TT who have described the lung of Heloderma have 
noticed this—to my eyes, very striking—peculiarity of the lung. 
Nor do the illustrations given by them show any signs of the 

* P.Z.S. 1890, p. 233. 
+ Deutsches Arch. f. d. Phys. 1818, Bd. iv. t P. Z. 8. 1861, p. 112. 
§ Zool. Jahrb. (Abth. f. Anat.) vii. 1894, p. 581, 
|| Zoe. cit. Taf. 31. figs. 16-18. 
§| Loe. cit. p. 581, fig. R, Taf. 32. figs. 19-21. 
** Especially fig. 20 of pl. 32 of his memoir. 
++ Shufeldt, loc. cit. p. 202; Stewart, P. Z.S. 1891, p. 118; Miller, “The Structure 
of the Lung,” Journ. Morph. vii. 1893, p. 170. 


eparterial bronchus in Heloderma. ‘The length of the bronchi 
and the complicated structure of the lungs themselves have, 
however, been remarked upon, and in these matters Heloderma 
also agrees with Varanus. The figure (text-fig. 16) shows the 
commencement of the lungs in Heloderma suspectwm as seen 
from the dorsal aspect. The long headward prolongation of 
each lung, not perhaps sufficiently emphasised in Dr. Shufeldt’s 
figure of these organs, is very reminiscent of Varanus. The 
rings of the trachea are, as correctly stated by Shufeldt, in- 
complete in the dorsal median line. The bronchi are described 
by the same writer as “unusually long”; but it is not clear from 

Text-fig. 16. 

A portion of the trachea, the bronchi, and the upper parts of the lungs of 
Heloderma, from the dorsal side. 

The left lung cut open to show the course of the bronchus within it. 

the description given, as Prof. Stewart * has pointed out, that this 
great length does not apply to the intrapulmonary portion of 
the bronchus. My own measurements of the length of the 
bronchus up to the place at which it enters the lung agree with 
those of Prof. Stewart in the case of Heloderma horridum. 1 
find this length, in fact, to be as nearly as possible 13 mm. ‘To 
be absolutely exact is impossible, on account of the pliability of 
the bronchial rings and interspaces. The bronchi of Heloderma 
are therefore shorter than those of Varanws (text-figs. 18 & 19, 

* P.Z.S. 1891, p. 120. 


pp. 65, 66), and not very much longer than those of a similarly- 
sized Iguana tuberculata. Generally speaking, it is undoubtedly 
correct to describe the Lacertilia as possessing short bronchi, to 
which rule, indeed, Varanus offers the only very marked exception. 
The relations of the bronchi to the lungs are not shown in 
Shufeldt’s figure *, where the heart obscures the same, and are 
wrongly shown in the figure of Millery. The latter author is 
wrong (unless, indeed, the lungs of Heloderma suspectum, examined 
by myself, are abnormal) in not indicating, in the figure referred to, 
a conspicuous branch of the bronchus developed equally on both 
sides of the body. When the trachea divides, the dorsal median 
fibrous wall lying between the disjunct ends of the tracheal 
semirings is continued down each bronchus. As Dr. Shufeldt has 
remarked, the calibre of each bronchus is not far short of that of 
the trachea itself. They are, in fact, particularly wide. The ends 
of the bronchial semirings are, of course, visible on either side of 
the median fibrous tract. ‘he bronchus approaches the lung and 
becomes adherent to its mesial side and runs down in contact with 
it for some distance until it finally enters the lung. At the point 
of contact the upper ends of the’ semirings, 7. e. those lying 
headwards, cease to be parallel with the lower ends and diverge 
headwards. The dorsal membranous space ceases, and the semirings 
in that section of the bronchus which is closely applied to the lung 
embrace the lung. There is, in fact, a branching of the bronchus, 
and this short branch may be seen to be lined by cartilaginous 
semirings for a short distance into the interior of the lung. This is 
not the case with the following apertures of communication between 
the bronchus and the lung. I cannot but think that this branch 
is comparable to that already referred to in Varanus. It 1s 
further not without importance to notice that this “ eparterial 
bronchus” in Heloderma is not serially comparable to the apertures 
which place the cavity of the bronchus into communication with 
the interior of the lung and which follow it. For the latter are 
more ventral in position, as is plainly to be seen in the accompanying 
figure (text-fig. 16). The ‘eparterial bronchus” is more dorsal 
and is, in fact, lateral with reference to the main stem of the 

The figure (text-fig. 16) which illustrates the branching of the 
bronchus before entering the lungs also shows on the left side 
the interior of the lung as seen when the bronchus is slit up after 
it has given off the branch referred to. I have thought it worth 
while to introduce this view of the lung of Heloderma, since the 
figure given by Miller ¢ does not appear to me to represent quite 
accurately the mode of communication between the interior of 
the bronchus and the lung-substance, nor does he indicate the 
adherence of the bronchus to the lung for a considerable distance 
before entrance. He does, however, illustrate the important fact 

* Toe. cit. pl. xvi. fig. 3. + Journ. Morph. 1898, pl. vil. fig. 5. 
t Loe. cit. pl. vii. fig. 5. 


that the bronchus traverses a considerable distance within the 
lung before it disappears. My own illustration will show that 
the semirings of the bronchus are complete for a considerable 
distance, and perfectly easily recognisable, since they show no 
particular differences from the semirings in the extrapulmonary 
region of the bronchus. The bronchus communicates with the 
lung by copious apertures, which are not situated in the region 
of the bronchus corresponding to the fibrous band which unites 
the tips of the semirings in the extrapulmonary region of the 
lung; these apertures would seem to be rather breaks in 
continuity of the semirings themselves. Their disposition is 
thus reminiscent of the way in which the rudimentary lung of 
certain Snakes arises from the bronchus. There is a simple hole 
in the bronchus which leads into the lung in the case of those 

In a paper communicated to this Society a good many years 
since * J described the complicated branching of the cystic duct 
and its anastomosis with the hepatic duct in Varanus salvator. 
T have since then discovered that the same network, comparable 
to that which is found so generally among the Ophidia, occurs 
also in V. gould, though it is in that species rather less developed 
than in V. salvator. Quite recently I have dissected out the bile- 
ducts in V. nloticws, of which species I have had the opportunity 
of examining several very small examples preserved in spirit. 
Thad one of these injected from the gall-bladder, and the injection 
(chrome-yellow rubbed up im olive-oil) ran readily along the 
branches of the cystic and hepatic ducts. The accompanying figure 
(text-fig. 17, p. 64) is fairly accurate (but I fear not absolutely 
so) as regards the network, which, as will be seen, is much like 
that of V. salvator, but perhaps not quite so complicated. More- 
over, when once the hepatic and cystic ducts have left the surface 
of the gall-bladder there are apparently no further anastomoses 
between them, as there are—though to a limited extent—in 
V. salvator. On the other hand, there are some species of Varanus 
in which there is no network formed by the bile-ducts on their 
emergence from the gall-bladder (as is also found among the 
Ophidia). To some of these I have referred in my communication 
just quoted. I have since carefully examined Varanus exanthe- 
maticus, and find that the cystic duct emerges as, and continues 
to be, a simple duct throughout. ‘The same is the case with 
V. griseus. It is not wise perhaps to generalise on these few 
data; but so far as the facts go they agree with an important 
external character by which the species referred to may be 
grouped. In JV. salvator, V. gouldit, and V. niloticus the nostril is 
a circular aperture, while in the other species mentioned it is 
obliquely placed and slit-like. 

In all the species of Varanus that have heen referred to in the 

* P.Z.S. 1888, p. 106. The illustrative figure (fig. 4, p. 105) has been copied in 
Gegenbaur’s ‘ Verg]. Anat. Wirbelth.’ 


present communication the pancreas is a solid body, as shown in 
the figure of that of V. niloticus (text-fig. 17). In all of them a 

Text-fig. 17. 


Pancreas, gall-bladder, &c. of Varanus niloticus, to show the course of 
the bile-ducts. : 

g-b. Gall-bladder; 4.d. Hepatic ducts; Z. Liver; O. Orifice of conjoined bile-ducts ; 
P. Pancreas; P.v. Portal vein. 

very slender splenic lobe arises near to the anterior end of the 
pancreas and passes to the spleen*. The species, however, show 

* In Varanus exanthematicus the end of the splenic not merely touches, but 
enwraps and is enwrapped by the spleen. This intimate relation between the 
spleen and the pancreas recalls a similar close association occasionally found among 
the Ophidia. 


certain differences among themselves in the position of the 
duodenum with reference to the pancreas. In J. niloticus the 
pancreas is separated by a considerable tract of mesentery from 
the duodenum, which results in the exposure of a long pancreatic 
duct. The same is the case with V. ocellatus. On the other hand, 
in V. griseus and V. exanthematicus the end of the pancreas 
touches the duodenum close to the point of entrance of the 
pancreatic and bile ducts, and there is therefore no great length of 

pancreatic duct. 
Text-fig. 18. 

Lungs of Varanus exanthematicus. 

Bronchi opened to show exit of “eparterial bronchi” close to pulmonary 
artery (P.a.). 

In his figures of the lung of several species of Varanus, Milani 
has pointed out certain differences which distinguish them. Thus 
the lungs of V. bengalensis are lobed externally in a fashion which 
is not to be found in the other species which are described. In 
examining, for the purposes of the present communication, the 
lungs of several species of Varanus, I have observed two small . 
points of difference between the lungs of certain species which are 
not referred to by Milani. Of these both involve an asymmetry 

Proc. Zoou. Soc.—1907, No. V. 5 


or symmetry of the lungs and the windpipe as the case may be, 
not, indeed, in point of length, which is a common feature among 
Lizards and one of the most obvious features in which they 
approach the Snakes. This asymmetry, when it occurs, affects 
the position of the “ eparterial bronchus” and the ventral forward 
projection of the lung between the bronchi. In Varanus griseus 
(text- fig. 19) the left lung, as well as the right lung, sends forward 
towards the bifurcation of the bronchi a thin diverticulum, which 
has an entire cavity not divided by any meshwork and seems to 
be comparatively unvascular. ‘This can readily be lifted up and is 
seen to be not attached to the bronchus of the lung of which it 
is an outgrowth. 

Text-fig. 19. 

Lungs of Varanus griseus. Details as in text-fig. 18. 

So far, therefore, the lungs are symmetrical. But the origin of 
the eparterial bronchus is not symmetrical. On the right side 
this branch arises from the bronchus at a distance of 27 mm. from 
the point of bifurcation of the bronchi; in the case of the left 


lung this distance was only 21 mm. <A second specimen showed 
precisely the same relations in all these points of structure. 
Contrasted with this (compare text-figs. 18, 19) are the different 
conditions observable in Varanus exanthematicus. In the latter 
species (text-fig. 18, p. 65) the two branches of the bronchi were 
exactly symmetrical and each was situate 33 mm. from the bifurca- 
tion of the bronchi. Only the right lung gave off a forwardly 
directed lobe situated on the inner side of its bronchus. There was 
nothing to correspond in the left lung. I have not been able to 
compare these conditions with those of many other species of 
Varanus. But in both JV. ocellatus and V. niloticus there was 
precisely the same asymmetry in the relative positions of the 
branch of the bronchus, which in all cases lies behind the aorta of 

its side. 

Summary of more important new Facts contained in this 

In view of the fact that very few genera and species of 
Lacertilia have been studied anatomically, it is a little difficult at 
present to differentiate between more and less important structural 
details as evidence of affinities between different genera. The 
following réswmé, therefore, will be necessarily only an attempt to 
lay stress upon what appear at present to be the more important 
new facts which I have set forth in this communication. 

(1) The pancreas in the Lacertilia, as already known, differs in 
different genera. I have added to the existing knowledge some 
new facts with regard to genera and species not examined by 
others. It appears from this that the chief variability in the 

ancreas consists in the presence or absence of a splenic lobe and 
in the relations of the latter to the spleen. The classificatory 
importance of the facts does not appear to be great ; since, though 
the Iguanoids, Zgwana and Liolemus, are like each other in the 
relations between the splenic lobe of the pancreas and the spleen, 
we find in Varanus and Chameleon differences between different 
species in these points. The pancreas is nearly always a compact 
gland; but not so in Zonurus. 

2) The variations in the structure of the viscera among the 
Chameleons concern principally the proportions between the two 
lobes of the liver, the form of the diverticula of the lungs and the 
absence or presence of these, the degree of pigmentation of 
the body-cavity, and the degree of coiling of the intestines. 

(3) The variations in the structure of the viscera in the 
different species of Varanus concern principally the presence or 
absence of a bile-duct network and certain minute differences in 
the lungs. It seems possible that those species with a round 
nostril are distinguished from those with an oblique slit-like 
nostril by the possession of this network. 

(4) The simplicity of structure which is often associated with 

small-sized forms as compared with their allies of larger size is 


well seen in the two small species of Chameleons, viz. Ch. pumilus 
and Ch. teniobronchus, where the lungs have no diverticula and 
the intestinal tract is nearly straight. 

(5) The very general presence of a gubernaculum cordis among 
the Lacertilia renders its absence in Zonurus a matter worthy of 

(6) The most important fact, perhaps, which I have been able 
to ascertain is the persistence in several genera of Lacertilia 
of considerable remains of the fourth visceral arch (second 
branchial). This is a fair-sized bar of cartilage which does not 
make any connection with the copula below. The existence of 
this arch has, however, been recorded in the adult Lacerta by the 
late W. K. Parker. 

(7) The two genera Varanus and Heloderma (which are quite 
remote from each other in some structural features) agree with 
each other in that each bronchus is adherent to its lung for 
some little distance before it enters it, and emits a short branch to 
the upper end of the lung before it becomes confluent with 
the lung. 

(8) It is interesting to note the double gastro - hepatic 
membrane in certain Scincide, which is associated with a 
correspondingly double set of gastro-hepatic veins, as distinctive 
of that family, though not universal. 

4, A List of Moths of the Family Pyrahde collected by 
A. E. Pratt in British New Guinea in 1902-3, with 

Descriptions of new Species. By Grorce H. Kenrick, 

[Received December 8, 1906. | 

(Plates III. & IV.*) 

This collection was made under circumstances mentioned by 
Mr. Pratt in his book ‘ Two Years among New Guinea Cannibals,’ 
published in 1905, and beyond the fact that most of the speci- 
mens were taken at light very little information can be given. 

The country in which the collections were made appears in 
some of its characters to resemble Darjiling: there are the same 
precipitous ridges with narrow valleys between, all with a back- 
ground of snowy mountains of great elevation, and everywhere 
there is much dense forest. The climate, with its abundant rain 
in the wet season and brisk air in the dry season, is also similar, 
while an abundant lepidopterous fauna completes the resemblance. 

Although most of the insects were taken at light, in most cases 
females were well represented and the condition of the insects is 
extremely good. 

* Hor explanation of the Plates, see p. 87. 

12 BS, USO, IPA. ih, 

EC. Knight delet lith, West,Newman chromo. 


124, SAO, IL INV. 

~ om SR a ae em Vers ta 
Sem i (0 Wn me) em 

West, Newman chromo. 

E.C. Knight Ii 








Subfam. SCH@NOBIIN. 
4, C. CACONALIS Swinh. 

MonopontTA, gen. n. 

Palpi porrect, more than length of head, parallel, clothed with 
hairs, but very thin; antenne simple; patagia shorter than thorax. 
Venation of fore wing: veins 2 and 3 before end of cell, 4 and 5 
from end of cell, 6 and 7 from end of cell, curved and nearly 
parallel, 9 and 10 from cell. Hind wing: cell rather short ; veins 
2 and 3 before end of cell, 4 and 5 from end of cell, 6 from upper 
angle, 7 anastomosing with 8 halfway between end of cell and 
end of wing; costa slightly curved; hind margin convex, rounded 
at angle. 

Type, J. passalis. 

5. MoNnoponta PASSALIS, sp. n. (Plate IV.) 

Head, legs, thorax, and patagia dark brown, abdomen rather 
paler. Fore wing pale brown inclined to purplish, an oblique 
dark line at one-fifth of the length of the wing, followed by a 
much darker portion outwardly bounded by a curved dark line 
deeply notched near angle of wing, this is followed by a pale 
ochreous shade. Hind wing pale ochreous at base, shading into 
darker towards the margin and having an indented dark line 
similar to that of the fore wing; fringes darker. 

Exp. 50 mm. 

Hab. Kebea, one specimen. 

Subfam. PHycrrin 2. 

6. PHycrra pinawa, sp. n. (Plate IIT.) 

Head, legs, palpi, and body ochreous, darker above. Fore 
wing marbled with purple, green and gold, the raised scales 
beyond the cell darker; a curved postmedian line dark margined 
with paler. Hind wing plain purplish grey. Fringes of fore wing 
purple ; fringes of hind wing paler. 

Exp. 24 mm. 

Hab. Dinawa, four specimens. 

7. EYIELLA FUSCALIS, sp. n. (Plate ITT.) 

Head, legs, palpi, and body brown with numerous pinkish hairs. 
Fore wing pinkish brown, a darker brown mark two-thirds along 
the inner margin edged on either side with paler; fringes dull 
pink. Hind wing smoky grey; fringes same colour. 

Exp. 28 mm. 

Hab. Dinawa and Kebea, four specimens. 

8. HypstpynA roBusta Moore. 


Subfam. EHPrPaASCHIINé. 
9. Macauua @NnocHRoa Hmpsn. 
10. M. prcta Warr. 
11. M. nycricHroaLis Hmpsn. 
12. M. semMINIVEA W1k. 
13. M. marearira Butl. 

14, M. UNIPUNCTALIS, sp. n. (Plate IIL.) 

Head, legs, palpi, and antennze ochreous, tarsi barred with 
darker. Fore wing reddish ochreous, with wavy antemedian and 
postmedian lines dark brown, between them is a black spot very 
near costa; the outer third of the wing is brown and there is a 
marginal row of black dots. Hind wing much paler, with similar 
marginal row of black dots and a faint curved median line. 
Fringes ample, shining brown. 

Exp. 28 mm. 

Hab. Dinawa, one specimen. 

15. M. CARADRINIFORMIS, sp. n. (Plate ITT.) 

Head, antenne, palpi, and legs ochreous, tarsi darker; thorax, 
patagia, and fringes all of the same colour. Fore wing ochreous, 
with curved and rounded postmedian line paler but outwardly 
margined with darker ; on the costa are three equidistant badly 
defined, slightly darker markings; from the middle of the base 
of the wing is a ragged darker streak, and this becomes stronger 
as it approaches the postmedian line near the end of which is 
a dagger-like marking. Hind wing uniformly ochreous. 

Exp. 32 mm. 

Hab. Dinawa and Kkeikei. 

16. M. pomauis, sp.n. (Plate III.) 

Head, thorax, palpi, and processes pale green, tibie barred 
with pale green and brown; eyes black with a network of buff ; 
abdomen green above, buff below. Fore wing apple-green with 
irregular markings of dark chestnut; a transverse band at one- 
third length of wing, an irregular divided blotch on inner margin 
reaching nearly to the angle, and a square-shaped blotch on costa 
near, but not at, the tip of the wing; a dark spot at end of cell. 
Hind wing pale brown ; fringes paler. 

Exp. 24-26 mm. 

Hab. Dinawa. 

17. M. cuRTULALIS, sp. n. (Plate ITT.) 

Head, antenne, palpi, and legs pinkish buff, tarsi ringed with 
darker. Fore wing wainscot-colour with a conspicuous chocolate 


tip; three dots on costa before this mark and a dot at end of 
cell. Hind wing pale clouded on margin with smoky. 

Exp. 23-26 mm. 

Hab. Dinawa. 

18. M. apicauis, sp. n. (Plate IIT.) 

Head, antennee, and palpi chestnut, thorax and abdomen the 
same with a few grey hairs; patagia paler. Fore wing; ground- 
colour chestnut, but the band, which is conspicuously white, is 
extended in some specimens in a tooth into the hind margin while 
on the other side it produces a basal blotch. Hind wing fuscous 
edged with darker. Fringes of both wings pale chestnut. 

Exp. 30-34 mm. 

Hab. Kebea, Dinawa, Babouni, Ekeikei. 

19. M. TENEBROSALIS, sp. n. 

Head, legs, palpi, and antennz ochreous; abdomen grey, with 
dark transverse band on second segment. Fore wing ochreous with 
basal and apical blotch; the band begins near the base with a dark 
interrupted line, it is twice as wide on costa as on inner margin 
and the exterior boundary is an oblique angulated dark line; the 
interior of the band is reddish ochreous with a dark dot at end 
of cell. Hind wing smoky. Fringes of both wings ochreous. 

Exp. 32-36 mm. 

Hab. Kebea, Ekeikei, Babouni. 

20. M. PERDENTALIS, sp. n. (Plate III.) 

Head, legs, palpi, and antenne pinkish ; thorax, patagia, and 
abdomen ochreous grey, with a slight chestnut band on second 
segment. Fore wing ochreous grey with dark lines and chestnut 
patches, basal blotch grey ; band very irregular, bounded on 
inner side by an oblique dark angulated line, on outer side by a 
strongly marked line, very narrow on inner margin but occupying 
about one-third of costa and mostly chestnut; beyond the band 
is a darker cloud towards apex ; there is also a dark central dot. 
Hing wing smoky with darker veins, rather paler towards the 
base. Fringes of both wings pale grey. 

Exp. 30 mm. 

Hab. Dinawa, Kebea. 

21. M. PORPHYREALIS, sp. n. (Plate IIL.) 

Head, legs, palpi, and antennz pinkish ochreous ; thorax grey 
to reddish. Fore wing mostly dark brown, with whitish and 
grey interrupted transverse bands; the median band is very 
indistinct but its outer boundary is marked by a paler narrow 
angulated band with a tooth in the middle; there is a row of 
subterminal dark spots. Hind wing straw-colour with darker 
margin. Fringes of both wings pale. 

Exp. 34-36 mm. 

Hab. Ekeikei, Kebea, Dinawa. 


All the species of Macalla enumerated above exhibit considerable 
colour variation and in most cases a certain amount of sexual varia- 
tionas well. It is quite possible that when they are better known 
some may prove to be varieties; on the other hand, it is also 
possible that some now treated as varieties may turn out to be 
veritable species. 


23. P. TRUNCALIS, sp.n. (Plate IIT.) 

Top of head, antennz, palpi, legs, and thorax pale buff with 
some black hairs; patagia black. Fore wing buff with a darker 
cloud near the angle and an interrupted subterminal double line; 
a subtriangular dark mark extends from near the base along the 
costa gradually thinning out; in the midst of this is the transverse 
brush of raised scales nearly black. Hind wing reddish with 
dense rufous hairs near body. Fringe of fore wing very dark ; 
fringe of hind wing reddish. Tibiz of hind legs clothed with 
rufous hairs. 

Exp. 38 mm. 

Hab, Dinawa. 

24, LOcASTRA CASTANEALIS, sp.n. (Plate IIT.) 

Head, legs, antenne, and palpi pale chestnut ; tarsi ringed 
with darker ; thorax and patagia chestnut, abdomen paler. Fore 
wing chestnut, the first third suffused with grey near the inner 
margin, after this comes a dark transverse angulated line forming 
the inner boundary of the band, the outer boundary of which is 
a curved pale transverse line; at the angle of the wing is a 
conspicuous grey elliptical blotch. Hind wing grey; fringes paler. 

Exp. 36 mm. 

Hab. Dinawa. 

27. S. prRAsSINA Warr. 
28. S. DIvITALIS Guen. 

29. S. FLAMMEALIS, sp.n. (Plate IIT.) 

Head and palpi greenish, antennze and legs pink, tarsi barred 
with darker, thorax and process greenish; body pinkish. Fore 
wing: ground-colour dark olive-brown; at one-third distance 
along the costa is a broad transverse band and at one half a 
distinct angulated but narrower band; these markings and a 
portion of the wing near the hind margin are of pale olive-green. 
Hind wing rich golden orange. Fringes of both wings orange- 

Exp. 26 mm. 

Hab. Ekeikei, one specimen. 


30. S. cornucaLis, sp. n. (Plate IIT.) 

Head, palpi, antenne, legs, and thorax rufous buff. Fore wing 
buff with darker and lighter lines and dots; on the costa a dark 
mark at base and another at one-third length of wing, smaller 
dots at one half and two-thirds, a cloud at the tip; a wavy 
subterminal line dark edged with lighter; a dark dot on inner 
margin near body. Hind wing buff, with a dark wavy line edged 
with lighter and dark central spot; fringes buff spotted with 

Exp. 22 mm. 

Hab. Dinawa. 

31. S. SUBVIRIDALIS, sp. n. (Plate ITI.) 

Head, legs, thorax, abdomen, palpi, and the long processes 
which extend to the back of the thorax, buff. Fore wing dull 
brown, with two transverse pale greenish bands parallel to hind 
margin, the first at one-half and the second at two-thirds length 
of wing; below the cell these bands run into a large patch of the 
same colour which continues along two-thirds of inner margin 
and projects a little beyond the second transverse band. Hind 
wing dull brown ; fringes same colour. 

Exp. 28 mm. 

Hab. Kebea, Dinawa, Babouni. 


33. O. POLYCHROALIS, sp. n. (Plate ITT.) 

Head, antenne, legs and thorax chestnut inclining to pink ; 
abdomen rather paler, with no band on second segment. Fore wing 
greenish with chestnut patches; basal blotch mingled olive and 
chestnut, with raised scales in longitudinal dark line, then a vertical 
line edged outwardly with white; the band is badly defined but 
there is an angulated thin line forming the external boundary ; 
the remainder of the wing is greenish white, with a chestnut 
apical patch. Hind wing pale buff. Fringes of both wings pale ; 
underside of costa of both wings broadly red. 

Exp. 30 mm. 

Hab. Dinawa. 

34. O, CHIONALIS, sp.n. (Plate III.) 

Palpi simple in both sexes, reaching vertex of head ; antennz of 
male ciliated and with a process consisting of an immense tuft of 
scales reaching over part of thorax; in the female this is very 

Head, palpi, antenne, and legs pinkish buff, legs ringed with paler 
at the joints; thorax, patagia, and process greenish; abdomen 
greyish ochreous. Fore wing: ground-colour dark olive-brown, 
basal patch with raised scales both light and dark followed by an 
oblique narrow white band, then follows the main band, in some 


specimens suffused with white and bounded by a curved paler 
line after which is a darker cloud. Hind wing fuscous; fringes 
of both wings paler. 

Exp. 38-39 mm. 

Hab. Ekeikei, Babouni, Kebea, Dinawa. 


Antenne of male ciliated, with process roughly scaled and 
reaching thorax ; female without process. 

Head, palpi, antenne, legs, and top of thorax ochreous; the 
male has a dark collar to thorax, absent in the female ; abdomen 
ochreous. Fore wing: ground-colour fuscous; a dark basal blotch 
occupies one-third of the wing; the band is bounded on the 
inner side by a narrow vertical white band, this is followed by 
pale ochreous with dark dots on the costa and is bounded out- 
wardly by a dark wavy interrupted line; the remainder of wing 
of the ground-colour with a row of subterminal triangular dots. 
Hind wind fuscous ; fringes paler. In some specimens the band 
is suffused with chestnut. 

Exp. 26-30 mm. 

Hab. Dinawa and Kebea. 

36. O. COLUMBALIS, sp. n. (Plate ITI.) 

Head, legs, antennze, thorax, and body dove-colour ; legs ringed 
with black. Fore wing dove-colour, on costa at base a triangular 
black patch, the apex of which reaches the inner margin as a 
thick black line and forms the inside vertical boundary of the 
band; the band itself is dove-colour, but there is a thin curved 
black line parallel with inside boundary and an irregular black 
patch on vein 2; the outer boundary of the band is a curved 
thick black line, and beyond this on the costa is an anteapical 
black patch. Hind wing dove-colour ; fringes paler. Underside 
smoky with whitish dots on costa. 

Distinct from seminivea in absence of white thorax and black 

Exp. 36 mm. 

Hab. Kebea, one specimen. 


39. KE. persicopa Meyr. 

41. C. atuensis Butl. 





Fore wing: vein 2 present; veins 3, 4,5 from lower end of 
cell; veins 6, 7 from upper end of cell; vein 7 parallel and close 
to 8 for one-third of its length, then turning down; vein 9 from 
middle of cell. Hind wing: veins 7 and 8anastomosing. Proboscis 
well developed ; palpi upturned reaching vertex of head, densely 
clothed with hairs ; antennz simple, three-quarters of the length 
of fore wing. Abdomen projecting for half its length beyond 
hind wing; legs long and slender, with small spurs on the tibie ; 
fore tarsi somewhat hairy. Fore wing long and narrow; hind 
wing subquadrate. 

42. 'T. TRIANGULALIS, sp. n. (Plate III.) 

Thorax, head, and palpi grey ; patagia whitish ; underside 
quite white; abdomen grey above, white beneath, tuft white; 
eyes very prominent. Fore wing dark smoky-grey shot with 
reddish; costa narrowly buff for two-thirds of wing; a small basal 
spot white, followed by three other white dots along the costa ; 
the inner margin with a narrow pale streak expanding into a 
quadrangular blotch at the angle. Hind wing white; a marginal 
dark band tapering from apex of wing and interrupted by a tooth 
in the middle. Fringes of both wings dark at apex and white at 

Exp. 36 mm. 

Hab. Dinawa. 

Subfam. PyRaLin#®. 
44, V. zemirA Cram. 

45. V. GRISEATA, Sp. n. (Plate IIT.) 

Head, thorax, and patagia pale orange, with four black dots 
across the front of thorax; abdomen black ringed with yellow; 
palpi black. Fore wing pale greenish grey ; veins near the hind 
margin tinged with black; a vertical metallic blue streak across 
the base of the wing and the space between this and the body 
dull orange, with one blue spot and a blue oblique streak below 
it; fringes same colour as fore wing. Hind wing black; fringes 

Exp. ¢ 51 mm., 2 66 mm. 

Hab. Dinawa. 

46. HypsopyGia postruAvA Himpsn. 

48. S. nigRopuncTata Hmpsn. 


Subfam. HypDRocAMPIN«A. 
52. A. ACROPERALIS Hmpsn. 
53. A. DIOPSALIS Hmpsn. 

55. A. LUNALIS, sp.n. (Plate III.) 

Head, thorax, and patagia buff; first segment of abdomen brown 
followed by a pale band, the remainder golden. Fore wing dark 
brown ; a pale brown stripe runs from the body and reaches the 
costa about midway; it is then continued as a fine line along 
the costa to the apex; along the inner margin is a whitish 
stripe with an angulated upward projection towards the middle 
of the wing; on the disc is a crescent-shaped white mark and 
this is followed by a white subterminal line with an orange band 
beyond ; fringes pale. Hind wing: the base for about one-third 
of the wing is white, the remainder orange; on the margin are 
four black dots, the two exterior having white spots above them ; 
in the middle of the wing are two black lines nearly parallel to 
hind margin. 

Exp. 20 mm. 

Hab. Dinawa. 

56. A. PURPUREALIS, sp. n. (Plate III.) 

Head, thorax, and abdomen purplish grey ; fore legs pale ringed 
with dark. Fore wing purplish grey with orange markings; a 
curved irregular band beginning on the inner margin and joining 
an orange band along the hind margin which is preceded by a 
silvery band; nearer the base is a dusky orange patch followed 
by a crescent of purple and then a long triangular mark of orange. 
Hind wing purplish grey at base, with a broad median band 
followed by a darker band, and finally by a narrow orange marginal 
band in the middle of which are seven silver-studded black spots ; 
fringes paler. 

Exp. 25 mm. 

Hab. Dinawa. 

57. A. BIPUNCTALIS, sp. n. (Plate IIT.) 

Head pale brown, thorax and first segment of abdomen dark 
brown, followed by a slightly paler band; legs pale ringed with 
dark brown. Fore wing dark brown; with a white streak fading 
into buff on the inner margin ; a white subterminal line followed 
by an orange band; fringes yellow. Hind wing: basal third 
white, remainder orange, with a short white band bordered with 
dark near the angle and a dark line parallel to the first but nearer 


to the base of wing; three black dots on the hind margin, two of 
which have white dots above them. 

Exp. 23 mm. 

Hab, Dinawa. 


Head, legs, palpi, and patagia white; thorax black above; 
antenne and abdomen dark brown. Fore wing: costa straight, 
hind margin nearly at right angles for half the width of wing, 
then slightly oblique to angle of wing; ground-colour white, with 
black border to hind margin and a parallel transverse narrow 
band at one-third of wing; from where this meets the inner 
margin is a narrow oblique red band to base of costa; from the 
lower end of black line on the hind margin is a double red oblique 
streak reaching costa and extending to middle of wing, becoming 
smoky on reaching the inner margin. Hind wing: base and hind 
margin white, followed by a smoky band, becoming red nearer 
the body and containing three white specks, slightly angulated. 
Fringes white. 

A very striking species both in form and colour. 

Exp. 24-30 mm. 

Hab. Kkeikei, Dinawa. 



61. B. MELANOPERAS Hmpsn. 

62. B. GuaucinaAuis Himpsn. 


66. D. HoRocHROA Meyr. 

67. D. ToRRIDALIS, sp. n. (Plate IIT.) 

Head and thorax fuscous; abdomen and posterior half of 
patagia pale buff, last three segments of abdomen fuscous; tuft 
buff; legs and underside white. Fore wing fuscous shot with 
purple, a dark central spot and a transverse dark mark at end of 
cell; beyond this is a curved thin dark line; fringes paler. Hind 
wing yellowish, the extreme hind margin darker, shading 
gradually into the colour of the wing and almost destitute of 
fringe in the male. 

Exp. 33 mm. 

Hab. Kebea. 

68. D. UNICOLORALIS, sp.n. (Plate IV.) 

Vertex of head pale, palpi and thorax dark grey; posterior 
end of patagia and abdomen pale; tuft black and a thin black 


lateral fringe. Fore wing olive-grey shot with purple, a minute 
black apical dot and fine black dots on hind margin; fringe pale. 
Hind wing: outer margin as fore wing; internal area straw- 

Exp. 36 mm. 

Hab. Dinawa, one specimen. 


Fore tarsi naked. 

Head, legs, antenne, tarsi, thorax, and abdomen dark brown ; 
tuft whitish orange. Fore wing dark brown with faint purplish 
tinge ; an antemedian, median, and postmedian angulated trans- 
verse line darker; a white dot at end of cell and another in the 
middle of cell, also a faint dot where the postmedian line reaches 
the costa. Hind wing with two angulated darker lines. Fringes 
dark with lighter patch at angle of wing. 

Exp. 30 mm. 

Hab. Kebea. 

Subfam. PyRAUSTIN®. 

71. P. vactiFERALIS W1k. 






77. A. FuMosA Himpsn. 

78. A. EUDOXANTHA Hmpsn. 


80. A. SEMIPICTALIS, sp. n. (Plate IV.) 

Head, thorax, and front of abdomen yellow spotted with orange, 
last three segments of abdomen purple; tuft orange. Fore wing: 
basal portion pale yellow flecked with orange, abruptly and 
obliquely terminated by a patch of rich brown shot with purple 
occupying the rest of the wing, in the middle of which is a red 
irregular spot ; the costa towards the apex is paler, and there is 
an obscure darker curved line in the middle of the wing. Hind 
wing similar to the fore wing, but without the red spot. Fringes 
conspicuously white with dark interruptions, giving the idea of a 
scalloped margin. 

Exp. 30 mm. 
fab. Dinawa. 



85. R. sceLatTatis Led. 

86. R. LINEALIS, sp. n. (Plate IV.) 

Head and thorax golden; palpi, antenne, and legs brown ; 
patagia darker; abdomen golden, but darker on the margin of 
each segment; tuft golden. Fore wing whitish with purple 
gloss and margins edged with gold, with numerous connected 
streaks and blotches of purplish brown. Hind wing similar, but 
the chief blotch is at the apex and a secondary blotch at angle. 
Fringes paler. 

Exp. 50 mm. 

Hab. Dinawa. 


89. S. MARMORATA Lucas. 

91. B. aprpawis Led. 

92. B. INVERTALIS Snell. 

93. B. AuRoTINCTALIS Hmpsn. 

95. PrmocRocIs ANIGRUSALIS W1k. 
96. P. copropasis Hmpsn. 

97. ULOPEZA CRUCIFERALIS, sp. n. (Plate IV.) 

Head, antennee, legs, and palpi bright ochreous; abdomen same 
colour in female, but lighter im male; tuft pale. Fore wing 
ochreous clouded with purple; three equidistant spots on eosta 
are produced in widening angulated bands until they meet a 
longitudinal band on inner margin; in the middle of the outer 
band is a projection reaching the hind margin. Hind wing 
yellow ; fringes same colour. 

Exp. 26 mm. 

Hab. Ekeikei, Mafalu. 

98. NosopPHORA DISPILALIS Himpsn. 
99. N. FuLvALIS Hmpsn. 

100. N. semrrrirauts Led. 

101. N. BARBATA Hmpsn. 



104. C. congropaTaLis W1k. 

105. C. cAsTANEALIS, sp. n. (Plate IV.) 

Head, thorax, and abdomen above dark grey, a white band on 
first segment of abdomen. Fore wing black at base followed bya 
white band for one-third of wing, this is edged with black and the 
tip of the wing is also black, between them the space is bright 
chestnut ; on the costa and extending to bottom of cell is a white 
spot and there is a white dot on the inner margin. Hind wing: 
basal half white, remainder chestnut; margin broadly darker. 
Fringes dark. 

Exp. 34 mm. 

Hab. Ekeikei, Kebea, Mafalu. 

106. C. oceLLauis, sp. n. (Plate IV.) 

This insect resembles Rhimphalea trogusalis W1k., but differs in 
the following particulars :—The spot in the cell is developed into 
a distinct ocellus with white centre, black ring, and yellow space 
with outer dark line, and it is much more vivid on the under- 
side; the postmedian line is double with a yellow centre, and 
there is a distinct parallel subterminal line, the hind margin 
being yellowish with a dark cloud in the centre. 

Sir G. Hampson considers that its right place is in the genus 

Exp. 27 mm. 

Hab. Ekeikei. 
107. FrnopEs XANTHALIS Hmpsn. 

108. TygPANODES RADIATA, sp. n. (Plate ITT.) 

Head, thorax, and tuft orange ; collar black; legs grey. Fore 
wing grey with whitish hyaline patches and streaks; a dark spot 
at base of costa followed by pale orange, then a transverse narrow 
black band, after this a triangular white patch from costa to 
inner margin, the nervures 1 and 2 showing on this; beyond is a 
large irregular white patch, the apex being broadly dark grey ; 
fringe greyish yellow. Hind wing dark with the nervures 
whitish and an oval white patch beyond the middle; fringe 
greyer than in the fore wing. ‘ 

Exp. 44 mm. 

Hab. Kebea. 

109. NEvVRINA PROcoPIA Cram. 

111. P. euypHopatis Wk. 

112. P. opscuratTa Moore. 

113. P. Basauticatis Led. 

114. P. oponrosticra Hmpsn. 


115. P. mMarcarira Butl. 

116. DicHocrocis xANTHOCYMA Hmpsn. 
117. D. Bvaxatis W1k. 

118. D. pUNCTIFERALIS Guen. 


120. D. PARDALIS, sp. n. (Plate IV.) 

Head and palpi orange; legs paler; tarsi with dark rings ; 
antenne dark; thorax ochreous, spotted with dark brown ; 
abdomen ochreous with darker cloud on back; tuft dark. Fore 
wing pale ochreous, with narrow dark basal band and two narrow 
equidistant bands crossing from costa to inner margin where they 
are wider or branched; a subterminal dark patch which only 
extends across half the wing. Hind wing ochreous slightly shot 
with purple, a dark subterminal line and a narrow dark trans- 
verse line across half the wing. Fringes golden. All the markings 

Exp. 30-40 mm. 
Hiab. Mafalu. 

121. D. BrmacuLatis, sp. n. (Plate IV.) 

Head, palpi, antenne, thorax, and abdomen above wainscot- 
colour; legs and underside pale and shining; patagia extending 
a little beyond thorax. Fore wing greenish buff; three yellow 
patches on costa passing into orange and bordered with brown, 
extending a little below middle of wing, the outer patch is made 
up of two elliptical yellow dots. Hind wing smoky, tinged with 
golden and spotted with darker underneath ; fringes paler. 

Exp. 31 mm. 
Hab, Kebea, Dinawa. 


125. D. ALTHEALIS WI1k. 

126. BotyoDrEs ASTALIS Guen. 

127. B. FuAviBasaLis Moore. 

129. S. cHRoMALIS WIk. 

130. S. cHALYBIFASCIA Hmpsn. 
131. S. pissrprrauis Led. 


133. 8. sonitucis Hmpsn. 

134. S. PERNITESCENS Swinh. 
Proc. Zoot. Soc.—1907, No. VI. 6 







. suRALIS Led. 





. IrysaLis WIk. 



. cmSALIS WI1k. 

. BICOLOR Swains. 


. INDICA Saund. 


. UMBRIA Hmpsn. 



AAA DAD a AO a Oam @ oD Pom @ 






. EXAULA Meyr. 


. EZGEALIS Swinh. 






[Jan. 15, 


170. G. QUADRISTIGMALIS, sp. n. (Plate IV.) 

Male with antenne slightly pectinated ; female quite simple. 

Head, legs, antenne, and palpi bright ochreous; patagia in 
male twice the length of the thorax; thorax and body ochreous ; 
tuft black. Fore wing: ground-colour ochreous with slight 
purple irridescence ; three equidistant dark purple bands on the 
costa are produced until they meet an irregular band of the same 
colour from inner margin along the middle of the wing; a faintly 
darker subterminal line. Hind wing divided sharply into three 
triangular divisions; those on the costa and inner margin 
ochreous, but the middle one is iridescent and has a darker 
transverse bar; on the hind margin are three double black dots 
followed by three single dots all surrounded by metallic scales. 

Exp. 32 mm. 

Hab. Ekeikei. 

171. G. pupicatis, sp. n. (Plate IV.) 

Head, thorax, antenne, and abdomen pale ochreous above, 
shining white below ; palpi with third joint brown; legs white ; 
sides of tuft black. Fore wing hyaline with golden and purple 
reflections ; a postmedian angulated transverse line and a faint 
mark at end of cell. Hind wing similar to fore wing. Fringes 

Exp. 30 mm. 

Hab. Kebea. 

172. G. PAUCILINEALIS, sp. n. (Plate IV.) 

Head, thorax, and abdomen pale with darker longitudinal lines ; 
underside, legs, and palpi paler; antenne brown. Fore wing 
crossed by narrow dark bands not parallel and each having a 
central yellowish line; the third and fourth touch at costa, 
beyond them is an irregular subterminal line broader at the 
apex; the spaces between the bands are shining white. Hind 
wing with only two bands, forming a triangle with apex at angle 
of wing; a darker mark on hind margin. 

Exp. 24 mm. 

Hab. Dinawa. 

173. G. MULTILINEALIS, sp. n. (Plate IV.) 

Very much like the above, but the bands are so much wider 
that the space left for the white ground-colour is so much reduced 
as to suggest a dark wing crossed by numerous white lines which 
are not parallel. The details can be better shown in a figure 
than described. 

Exp. 31 mm. 

Hab. Dinawa, Ekeikei. 

174. G. PERSPICUALIS, sp.n. (Plate IV.) 

Head, top of thorax, and abdomen chestnut; patagia, sides of 
abdomen, and underside pure white; tuft black. Fore wing 


golden chestnut, with hyaline white blotches surrounded by 
darker lines; a basal triangular spot ; an antemedian oval spot; 
a postmedian ovate spot, three small dots between these two; an 
apical spot with a line of dots below reaching to inner margin ; 
two half spots along inner margin; fringes brown, white at 
angle. Hind wing hyaline with marginal dark band intersected 
by row of chestnut spots; fringes brown, spotted with white, 

Exp. 32 mm, 

Hab. Dinawa. 

175. G. EXQUISITALIS, sp. n. (Plate IV.) 

Head, palpi, and thorax brown ; abdomen in the male grey, in 
the female yellow; tuft black. Fore wing chestnut faintly shot 
with blue and purple; the spots hyaline and yellowish in the male 
but white in the female ; a streak from costa at base sloping to 
middle of wing in hind margin; an antemedian triangular costal 
spot with long tapering end; a broad postmedian costal spot 
extending almost to hind margin, with a darker outside margin 
and slight projection into the spot from outside; two very faint 
purple, parallel, subterminal lines; fringes golden. Hind wing 
pale straw-colour ; in the middle of the hind margin are a few 
black dots set in bright golden scales; fringes golden. In the 
male the hind wing is dark grey and the fore wing darker. 

Exp. 32 mm. 

Hab. Kebea. 

176. G. BRUNNEOMARGINALIS, sp. n. (Plate IV.) 

Head, thorax, and palpi green; antenne dark; abdomen green, 
but silvery beneath. . Fore wing apple-green with yellowish costa 
and margins; one black dot at end of cell and one in the middle. 
Hind wing green, one large black dot at end of cell, margins 
having a scorched appearance fading into white nearer the body. 
Fringes smoky, but inclined to chestnut at angle. 

Exp. 32 mm. 

Hab. Kebea. 

177. G. LACERITALIS, sp. n. (Plate IV.) 

Head, thorax, and palpi apple-green ; antennze and last joint of 
palpi brown; legs white, but tibie of fore legs with brown hairs ; 
abdomen green above, white at sides and bluish white beneath ; 
tuft black with white centre. Fore wing apple-green; hind 
margin distinctly and irregularly scalloped, edged with brown as 
if scorched ; one black dot in middle and one at end of cell. 
Hind wing apple-green, scalloped and scorched like fore wing; 
one black dot at end of cell; inner margin white and hyaline 
with fringe of same; other fringes brown. 

Exp. 44 mm. 

Hab. Ekeikei, Kebea, Dinawa, Mafalu. 


178. Pygospiua Bivirratis WIk. 

179, P. Mareinauis, sp.n. (Plate IV.) 

Head purplish grey; palpi, thorax, and abdomen dark grey 
above, shining white below; legs white; patagia grey shot with 
purple. Fore wing grey shot with purple; a white patch on vein 
below costa at two-thirds length of wing from base, 2 mm. wide, 
ending in three white dots towards inner margin; a second white 
patch between base and first patch; between this and angle of 
wing two smaller patches and one still smaller on inner margin ; 
fringes grey. Hind wing dark grey shot with purple, discal 
portion semitransparent with central spot and serrated dark band; 
fringes white. 

Exp. 45 mm. 

Hab. Mafalu. 

180. P. IMPERIALIS, sp. n. (Plate IV.) 

Head, thorax, and abdomen grey, striped with white; palpi 
black, first and second joints white; underside white. Fore 
wing dark grey shot with rich purple; a round white spot near 
base surrounded by four white dots; two median, two postmedian, 
and two smaller anteterminal white blotches. Hind wing: 
ground-colour as fore wing, one long triangular basal blotch, five 
other blotches on disc. Fringes dark, flecked with white at angle 
of wing. 

Exp. 36 mm. 

Hab, Ekeikei, Kebea, 

181. H&orria viressormpEs Moore. 

182. H. pominauis Led. 

183. PotyrHuiprA MACRALIS Moore. 

184. P. pivaricata Moore. 


186. Merra@a NEBULALIS Wlk. 

187. Leucrnopes apicaLts Hmpsn. 

188. SAMEODES TRISTALIS, sp. n. (Plate IV.) 

Head, legs, antennee, and palpi very dark grey; thorax slightly 
paler; abdomen with first three segments paler. Fore wing dark 
grey with a V-shaped white transverse mark having its apex on 
the inner margin ; near the outer margin of this mark is a thin 
angulated dark line. Hind wing dark with irregular transverse 
paler band ; fringes dark, spotted with white. 

Exp. 24 mm. 

Hab. Dinawa. 

189. Merrocrena stainronr Led. 

&6. MR. G. H. KENRICK ON PYRALID# [Jan. 15, 

192, A. EUcosmA Turner. 

193. A. I1GNEALIS WI1k. 


195, I. mrntauis Warr. 

196. Maruca TESTULALIS Geyr. 

197, PacHyNoa THOOSALIS W1]k. 

198. P. sprLosomorpEs Moore. 

200. P. PALLIDALIS Hmpsn. 


202. BaoraRCHA HYALINALIS Hmpsn. 
203. B. timpata Butl. 


205. C. ALBIPUNCTALIS, sp. n. (Plate IV.) 

Palpi dark purplish brown ; head orange; thorax and patagia 
purplish brown ; abdomen above fuscous; thorax and abdomen 
below, legs and underside of palpi white. Fore wing dark purplish 
brown, with a subtriangular patch from the middle of the costa to 
middle of the wing orange, paler inside and with a minute orange 
dot in the middle; hind margin broadly orange with a row of 
subterminal dots. Hind wing white on costa, the middle of the 
wing occupied by a triangular purplish patch, the apex reaching 
the these of the wing ; between this and the inner margin 1s walnihe 
the hind margin broadly orange and fringe orange; in the centre 
of the patch are two minute white dots edged with dark. 

Exp. 44 mm. 

Hab. Dinawa. 

207, P. nypstusALis WIk. 

209. P. BRACTEALIS, sp.n. (Plate IV.) 

Head, thorax, abdomen, and legs pale ochreous, inclining to 
reddish above but shining white below, very noticeable on tibia ; 
palpi with third joint chestnut. Fore wing golden yellow; costa 
margined with reddish; five equidistant, thin, angulated, some- 
what interrupted transverse lines, Hind wing the same, with 
three faint lines. Fringes pale. 

Exp. 30 mm. 

Hab, Dinawa, Ekeikei. 

210. Pyrausta CELATALIS WIk. 
211. P. ERtoprsatis W1k. 
212. P. ceapEsaLis WIk. 

213. P. rriricatis, sp. n. (Plate IIT.) 

Head and thorax orange; patagia darker; abdomen paler with 
chestnut markings along the crest; palpi and antenne dark 
brown; underside and legs whitish; tarsi ringed with darker. 
Fore wing orange with dark margin on the costa, dark nervures, 
and the cell margined with darker ; an oblique line, dark brown, 
passes through the cell and reaches the hind margin; a second 
oblique dark line reaches from the middle of the hind margin to 
near the apex. Hind wing straw-colour, margined with darker. 
Fringes smoky with white edges. 

Exp. 26 mm. 

Hab. Dinawa. 


215, P. puRPURALIS WI]k. 

216. OmpHisa INGENS Hmpsn. 

In concluding this list I desire to tender my most hearty thanks 
to all those who have assisted in determining the species and 
especially to Sir G. Hampson, Col. Swinhoe, and Dr. Dixey, who 

kindly allowed me to see the Walker types in the Hope Museum 
at Oxford. 


PratE Il. Prare IV. 
No. No. 

6. Phycita dinawa, p. 69. 5. Monodonta passalis, p. 69. 

7. Htiella fuscalis, p. 69. 58. Parthenodes rectangulalis, p. 77. 
14. Macalla unipunctalis, p. 70. 68. Dracenura unicoloralis, p. 77. 
15. na caradriniformis, p. 70. 69. Piletocera inconspicualis, p. 78. 
16. “ pomalis, p. 70. 80. Agrotera semipictalis, p. 78. 

17. = curtulalis, p. 70. 86. Rhimphalea lineatis, p. 79. 

18. i apicalis, p. 71. 97. Ulopeza cruciferalis, p. 79. 

20. 33 perdentalis, p. 71. 105. Caprinia castanealis, p. 80. 

21. - porphyrealis, p. 71. 106. 45 ocellalis, p. 80. 

23. Polylophota truncalis, p. 72. 120. Dichocrocis pardalis, p. 81. 

24. Locastra castanealis, p. 72. 121. > bimaculalis, p. 81. 

29. Stericta flammealis, p. 72. 170. Glyphodes quadristigmalis, p. 83. 

30. 3 cornucalis, p. 73. iil . pudicalis, p. 83. 

31. cs subviridatis, p. 73. 172. is paucilinealis, p. 83. 

33. Orthaga polychroalis, p. 73. 173. oo multilinealis, p. 83. 

34, es chionalis, p. 73. 174. = perspicualis, p. 83. 

35. S fuscofascialis, p. 74. 175. s exquisitalis, p. 84. 

36. D columbalis, p. 74. 176. i brunneomarginalis, 

42. Tipuliforma triangulalis, p. 75. p. 84. 

45. Vitessa griseata, p. 75. 177. 53 laceritalis, p. 84. 

55. Aulacodes lunalis, p. 76. 179. Pygospila marginalis, p. 85. 

56. D purpurealis, p. 76. 180. 35 imperialis, p. 85. 

57. % bipunctalis, p. 76. 188. Sameodes tristalis, p. 85. 

67. Dracenura torridalis, p. 77. 205. Calamachrous albipunctalis, p. 86. 
108. Tyspanodes radiata, p. 80. 209. Pionea bractealis, p. 86. 

213. Pyrausta triticalis, p. 87. 

88 DR. E. A. GOELDI ON MARMOSET [Jan. 15, 

5. On some new and insufficiently known Species of Marmoset 
Monkeys from the Amazonian Region. By Prof. {Dyes 
Bur A. Gorupt, C.M.Z.S., Director of the Para Museum. 

[Received November 23, 1906. | 
(Text-figures 20-23.) 

On the occasion of the Sixth International Zoological Congress, 
held at Berne (Switzerland) in August 1904, I presented a paper, 
‘Nova zoologica from the Amazonian Region, dealing especially 
with new Vertebrates,” in which I discussed at some length new 
and little-known representatives of the family of Callitrichidee 
(Hapalide, auctorum*) from the Upper Amazon, especially from 
the Rio Purtis, as follows:—(1) A species of J/idas whose close 
relationship with Midas rufiventer Gray I recognised at first 
sight, and to which later on, after comparing it with the 
type-specimen of this latter in the British Museum in London 
(at the time of my visit to the International Ornithological 
Congress in July 1905), I decided to give the name of Midas 
griseovertex. (2) A second species of J/idus, evidently related to 
M. labiatus Geoftr. (1812), MW. illigert (id. 1845), and M. weddelli 
(id. 1849), and having as its most characteristic distinguishing 
feature an enormously long white moustache, which afterwards, 
on the same occasion, I decided to name J/idas imperator. 
(3) Midas pileatus, described by Geoffroy in 1848 from the original 
specimen from the Rio Javary, kept in the Paris Museum, and 
until recently not represented in any other Museum, so far as 
could be judged from current zoological literature. I was then 
able to show a splendid pair of this very rare species. (4) Midas 
fuscicollis, described so long ago as 1823 by Spix, but only from 
immature specimens, the habitat of the adult animal of both sexes 
having escaped notice, as it appears, in a surprising manner until 
1904. (5) Midas mystax Spix, originally described from the Rio 
Solimoés, represented in my exhibited collection by a very dark 
specimen from the Rio Jurua. 

One of the essential conditions for arriving at certainty in my 
conclusions, as above suggested, was the careful examination of 
certain individuals kept as type-specimens in the British Museum. 
The purpose of the present article is principally to record the 
result of my investigation in this respect, which proves to be most 
interesting and instructive. To say it at once: the most remark- 
able and unexpected discovery was the fact, that of the two 
specimens of Midas rufiventer, type and quasi co-type, on the 
shelves in the British Museum, the latter does not coincide with 
the former, but, on the contrary, should constitute the type- 
specimen of a new, overlooked, and undescribed species of the 
genus JJidas. 

* See Thomas, Ann. Mag. N. H. (7) vol. xii. (1903). 


Minas THOMAST, sp. n. (1905). 

[Labelled in the British Museum: “ Midas rufiventer, 3, 
‘The red-bellied Tamarin, Upper Amazonia. W. Bates. 
Exp. 1857, Tunantins, north side of Amazons.” | 

Dorsau Aspect.—Anterior half of a dark, deep blackish-brown 
colour (somewhat like the half-grown J/, fuscicollis), extending 
from behind the ears as far as the middle of the back. Hinder 
half of the same brown, the tips of the hairs being however 
mottled with a lighter greyish-brown, but in a much less pro- 
nounced degree than in the typical JL. rufiventer. 

Head (text-fig. 20) black, with an insignificant median small 
light spot beginning just between the ears, behind the vertex ; 
a still less noticeable lightish dash, consisting only of a few 
greyish hairs, almost forming an anterior continuation, is per- 
ceptible in the same median line just in front of the former. 

Details of the region of mouth and nose as in the type of 
M. rufiventer. 

Text-fig. 20. 

Back view of head of Midas thomasi. 

Arms dark blackish-brown, passing into a true black towards 
the hands, not being, however, a brilliant black as in the 
M. rufiventer type. 

Legs. Yixterior side of thighs of the same colour as the posterior 
half of the back, turning to black toward the feet. 

Tail black. 

VENTRAL ASPECT.—Anteriorly a cross-zone of light reddish- 
yellow (viz., inside of arms and a band across the breast as wide 
as the insertions of thearms). Rest from there backwards reddish 
rust-yellow (lighter than in the new species MV. griseovertex G., 
and much lighter than in J/. rufiventer type). 

The light under side is abruptly separated in the region of the 
neck and breast from the black of the neck and head. (This 
light abdominal colour cannot be attributed to fading of an old 

90 DR. E, A. GOELDI ON MARMOSET  [Jan. 15, 

Museum specimen, perhaps unduly exposed to light on the 
shelves, because in that case such change would certainly be even 
more noticeable in the colouring of the back, which, although 
more exposed, still holds however its colour fast ; consequently 
the impression is produced of a decidedly distinguishing character 
of specific rank.) The light abdominal colour extends to the 
under side of the insertion of the tail, as in J. rufiventer type 
and in the other new species, JZ. griseovertea. 

On separating the fur the deeper portion is of the dark 
colouring, which is lighter in the anterior than in the posterior 
half of the body, and thus corresponding perfectly with the scale 
of colouring of the external aspect of the fur. 

The only existing specimen (stuffed) dates, as noted above in 
the copy of the label, from Tunantins, Upper Amazon, where it 
had been collected in 1857 by Henry W. Bates. 

Let us now proceed to the description of the type-specimen 
of the true Midas rufiventer Gray, m the British Museum, in 
order to relieve once for all the deplorably embarrassing situation 
caused by the very defective descriptions given up to the present 


[‘‘ Brit. Mus. Reg. Type (skin). Locality : 
South America. Purchased of Argent. ¢ (skull).”] 

I must state at the outset, that I was informed by Mr. Oldfield 
Thomas that the specimen had been obtained from a dealer, a 
fact which accounts for the vagueness of the locality. 

In general the results of my examination coincide with the 
description given by. Gray in his ‘Catalogue of Monkeys, ete.’ 
1870, p. 66, of this original specimen, only that it is too brief. 

DorsaL ASPECT.—Predominating colour of the back a dark 
blackish-brown. Hairs light greyish for the apical + of their 
length. Anterior part of back less greyish, posterior part. of 
back more so. The fur down to the roots dark brown (not 
light greyish-brown). 

Head (text-fig. 21). General colouring of the head a deep 
black, not blackish-brown. Mouth bordered with sparse whitish 
hairs. Central zone between and beneath the nostrils sparsely 
covered with lightish grey hairs (not brilliantly white). 

A spear-shaped dark rust-red median stripe extending from the 
vertex between the ears downward to the insertion of the nose, 
the rust-red beg of the same intensity as the colour of the whole 
abdominal side. From the vertex backward, in form of a double 
ploughshare, over the occiput down to the neck spreads out 
a patch of a greyish-brown colour, which combined with the lance- 
shaped frontal marking forms a comparatively broad triangle with 
a tendency to lateral posterior development. 

Arms. Outer side deep dark brown (like the head), slightly 
lighter on the upper arm. 


Legs. Outer side of the same colour as the posterior part of 
the back. 

VENTRAL ASPECT.—General colour a deep dark chestnut reddish- 
brown, considerably darker than in the Purus species, J/. griseo- 

Tail very long, dark blackish-brown, turning blacker toward 
the end. 

Text-fig. 21. 

Back view of head of IWidas rufiventer. 

So far as I know the only figure as yet published of a specimen 
of Marmoset Monkey, closely. resembling this ty pe-specimen of 
Midas rufiventer in the British Museum, is that in the supple- 
mentary plate 36 of the work of Reichenbach, under the name 
of “ Midas erythrogaster Natterer, Mus. Vindob.,” made, as stated 
on p. 14, from an original sketch by a Mr. T. E. Zimmermann 
of a skin collected by Natterer and preserved in the Vienna 
Museum. With no little surprise I discover, however, that 
Pelzeln, the excellent monographer of the Brazilian mammals 
collected by Natterer, enumerates this very specimen, and also 
the figure in Reichenbach, as synonyms under the head of J/das 
labiatus Geoffroy, a procedure which seems to me more than 
doubtful, not to say flatly erroneous. In this figure, though 
worthy of criticism in some respects (the most salient defect 
being the substitution of the delicate mottling by rough. daubs of 
‘ieee on a greyish ground), one essential feature nevertheless 
stands out clearly, namely the rust-coloured frontal spot, coinciding 
perfectly with the abdominal colour. 

I have never been able to consult either the full description, 
or any figure, or any other useful information concerning the 
M, elegantulus Slack, Proc. Academy of Natural Sciences of 
Philadelphia, 1861, p. 463, always quoted as a synonym of 
M. rufiventer Gray. 


Having now finished these explorations of the treasures stored 
up in the British Museum as an essential preliminary and 
solid basis for the discussion of the respective questions that arise, 
I will proceed to the detailed description of the two new species 
of Marmoset Monkeys of the Purts Region, mentioned at the 
beginning of the present article. - 


In general aspect closely similar to I. rufiventer Gray and 
M. thomasi Goeldi, in the sharp contrast between the bright 
rusty-coloured abdominal side and the dark colouring of the 
dorsal aspect, but distinguishable at a glance by the greyish- 
white, good-sized rounded frontal patch. 

Among the collections made in the Purts and Acre Regions 
by our Museum Expeditions (1903-1904) there are seven indi- 
viduals of this most interesting and well characterised species of 
Marmoset Monkey (skins, skulls of all and trunks of some), 
4 being malesand 3 females. One mounted pair (¢ @ ) remained 
in the Museum at Berne, labelled as above; three mounted indi- 
viduals (¢, 2, and a half-grown young one) are kept in the Para 
Museum ; and the remaining pair of skins (¢ @ ) are intended for 
the British Museum. 

Dorsau Aspect.—General colour deep blackish-brown, excepting 
head, hands, feet, and tail, which are positively black. This 
colour remains pure from the nape of the neck backward for 
one-third of the length of the back. The hinder two-thirds of 
the back shows a mottling due to the light greyish tips of the 
hairs, which terminal points measure about + of the entire length 
of the hair and are slightly longer toward the hips; the intensity 
of the mottling increasing gradually backwards, being most pro- 
nounced in the sacral region, presenting even a whitish appearance, 
when seen from certain oblique directions. 

On separating the fur in the region of the shoulders, the 
impression of colour is in general the same as the exterior, that 
is deep blackish-brown, excepting that the lower third of the 
hairs towards the roots forms a slightly lighter zone, especially 
laterally to the median line. Making the same examination in 
the region of the hips, the light zone is scarcely apparent, the fur 
being of the general colour almost down to the roots. 

Head. General colour sooty-black. A narrow zone of whitish 
hairs bordering the whole extent of the mouth. In the region 
of the upper lip the white zone rises in a very conspicuous 
triangular zone, with its broad base resting on the circular white 
band of the mouth. One very noteworthy feature of this 
triangular zone is that the lateral oblique lines cross exactly 
the middle of the nostrils, so that the exterior half of each 
falls in the blackish region, the interior half in the whitish 

At the vertex, from a line connecting the anterior borders 
of the ears, commences a whitish patch (text-fig. 22), occupying 


about 3 to 2 of the width between the ears and widening slightly 
backwards, but in no specimen acquiring the double ploughshare- 
shape of the rust-coloured patch of the Midas rufiventer type. In 
young individuals this patch is rather more greyish, while in 
more aged specimens it becomes nearly white. In length it 
extends as far as the posterior border of the occipital region. 

Arms. Outer side from shoulder to elbow of the same sooty 
blackish-brown as the whole shoulder region; fore-arms and 
hands decidedly black. 

Legs. Outer side of the same intensely mottled colour as the 
entire hip-region as far as to the foot; the foot itself of the same 
black as the hand. 

Back view of head of Widas griseovertex. 

VENTRAL ASPECT.— General colour light reddish rusty, decidedly 
lighter than in Midas rufiventer. It embraces the whole area 
from the middle of the. throat, breast, and belly, inner side of arms 
and legs, and a short distance beyond the insertion of the tail on 
its under side (some 5 cm.). On the flanks the dorsal and ventral 
colours are very abruptly separated, it being noteworthy that the 
rusty-coloured hairs of the abdominal side are only about half as 
long as the adjacent dark-coloured hairs of the dorsal part. 

Tail long, exceeding the length of the body by that of the head. 
Colour the same black as of the fore-arms and hands, but here 
and there with rows of hairs with light brownish tips, which may 
be interpreted probably as the vestiges of an annular arrangement. 


In general aspect, especially in colour, somewhat similar to 
M. labiatus Geoftr., MW. uligert, and M. weddellii, but easily dis- 
tinguished by the immense white moustache, which becomes 

94 DR. E. A. GOELDI ON MARMOSET [Jan. 15, 

phenomenal especially in the adult male. (Text-fig. 23 represents 
the head of an old male individual.) 

Ow: Purts collections embrace five skins (and skulls) (395 ¢, 
2 2 2), two from the Rio Acre and three from the upper Rio 
Puriis. <A family of three individuals (¢ 2 adults, and a young 
one) are already mounted in the Par&é Museum; the two 
remaining (¢, @) skins, of half-grown specimens, are destined 
to be sent to the Museums of Berne and London respectively. 

This species is distinguished by a more or less pronounced 
reddish greyish-brown general colour of the dorsal side, light rust- 
coloured abdominal side, dark feet, hands, and head, excepting 
the white cireumbuccal zone and the skull-cap, which tend to 

become grey with age. 

Head of Midas imperator. 

Dorsau aspect.—The reddish-rust colour is more conspicuous in 
the above mentioned old male, and is already noticeable in the 
two young males (one stuffed in the Para Museum and one skin). 
General colour as above stated ; it may be noted, however, that the 
greyish tinge, in the same way, is more pronounced in the female. 

On separating the fur nearly every one of the five specimens 
presents a slightly different appearance. In the two adult stuffed 
specimens in the Para Museum the lower half is uniformly dark ; 
the outer half shows alternately a light zone, then a dark one, 
followed by a light one and a dark one, the terminal zone being 
again lighter, making five zones of equal breadth in the outer half 
and giving the general impression of two equal pairs of light and 
dark zones. 


Examining in the same manner the two skins of half-grown 
individuals (2, ¢), we see immediately, that while the lower 
half is dark as above, the outer half shows alternately a light 
zone, then a dark one, and finally the terminal lighter one, giving 
as a total result the impression of only one complete pair of 
light and dark zones. The female skin presents nevertheless a 
peculiarity in the circumstance that near the roots of the hairs 
there is an unmistakable trace of a basal light zone—a peculiarity 
of this one individual among all the five known. 

Proceeding now to the examination of the same details in 
the quite young stuffed specimen of the mounted family in the 
Para Museum, the general impression is of only one well-marked 
pair of dark and light zones, from within outwards, while on closer 
inspection the second outer paix noticed in the half-grown speci- 
mens is unquestionably recognisable. 

Recapitulating these facts, the general conclusion is that the 
number of light and dark zones tends to increase with age. We 
have here one more useful hint, that in the discrimination of species 
prudence is required in judging the value to be attributed to 
such variations in colour, which in the present case can be fairly 
proved to be due to differences of age and sex. 

Head. All parts of the full front view of the face and direct 
lateral view are sooty-black in the adult female, the quite young 
individual (g) and the two skins (¢ 2), while the old male 
shows the face somewhat mixed up with greyish-brown hairs. 
The circumbuccal zone, however, is again an exception from 
this general colour, being saliently pure white. This zone is 
relatively broader than in any of the preceding species, including 
the entire nostrils in the old male, while in the other four speci- 
mens the border-line of the white colour curves below the nostrils, 
arising again on the ridge of the nose at about one-third of its length. 
The same white colour characterises the phenomenal moustache, 
which already in infant specimens is of a greater length than in 
any other of the moustached species known to me. The old 
male (text-fig. 23), however, beats the world’s record, the longest 
hairs attaining a length of no less than 57 to 60 mm.! 

My collectors inform me that the moustache is worn in life not 
straight, but with the modern upward twist. The two oldest 
specimens show the strange strong development of the white hairs 
of the circumbuccal zone extending even over the whole area of 
the lower jaw, not including the chin and inferior side of the jaw. 
As a result, the old males appear bearded as well as moustached. 

In the three younger individuals the crown of the head is 
brownish-black, with a tendency towards forming the already 
familiar favourite lighter-coloured patch of a pale brownish shade. 
In the adult female the patch is diamond-shaped, almost dirty 
greyish-white, and therefore very conspicuous. In the old male 
a pronounced greyish cast spreads all over the top of the head, 
without forming any distinct patch. 

Arms. Outer side of upper arm the same colour as the back, 

96 DR. E, A. GOELDI ON MARMOSEL | Jan. 15, 

but a little lighter in all specimens. Fore-arms and hands 
gradually darker towards the extremity. This is especially the 
ease in the three young specimens and the old female, while the 
male shows on the outer side of the fore-arm and hands the same 
dark greyish cast as noted above in the description of his face. 

Legs. Outer side of thigh the same colour as the back, but as a 
rule a little lighter, especially m the old female. The outer 
side of the foot is of the same dark colour as the hand. 

VENTRAL ASPECT.—The colour forms a marked contrast with 
that of the dorsal aspect, by having a pronounced tendency towards 
a rusty red, especially in the younger specimens. This rusty-red 
shade covers the whole under side, from the throat backward in- 
cluding the inner side of the arms and legs and no small extent of 
under side of tail. The same colour tends to form regions or 
patches of a deeper shade, one between the arms, covering the chest, 
and another between the legs, covering the abdomen and sur- 
rounding the anus. The chest-patch, especially in the old female, 
looks as if soiled by dried blood. 

Tail. The tail is long, nearly half as long again as the body 
(the average length of four specimens measured being as 11 : 8). 
In the younger specimens the dorsal side nearly from the inser- 
tion to the end is of the same dark colour as the feet, while the 
lower side shows a gradually narrowing line of the above mentioned 
deep rusty red fading toward the extremity. In the adult 
specimens the colour of the tail tends to become uniform in its 
whole circumference. In the old female the upper side is darker 
than the lower at least for two-thirds of its length, the terminal 
third being of a dark grey, due to the black roots and the ght tips 
of the hairs. The under side for about half its length has the general 
hight rusty-red colour of the ventral aspect, the terminal half 
being alike on the dorsal and ventral side, a dark grey. 

Most aberrant is the colouring of the tail in the old male. 
From the very insertion the bright rusty-red colour predominates 
in its whole circumference throughout its entire length with the 
exception of a dark terminal tuft. The rusty red is most 
noticeable to a certain extent (one-eighth) from the insertion 
backward. Approaching the end the colour is more mixed with 
dark, owing to the greater extent of black at the roots of the 
individual hairs. <A’ darker isolated patch exists also on the 
second eighth. 


This species, it is true, was long ago described, having been 
introduced into science in 1823 by the Bavarian explorer J. von 
Spix from individuals obtamed on the Rio Javary, but it is 
evident from our much richer material (eight specimens, 6 ¢ ¢ 
and 2 2 2), collected on the Rio Purtis, that the pelage of 
the full-grown animal has never been properly described and 
figured. Referring to a comparison of the two figures 3 (adult 
coat) and 3.@ (juvenile coat) of my original coloured Marmoset- 


Monkey plate, presented to the Zoological Congress at Berne and 
now being reproduced for a separate publication, it is especially 
noticeable that the adult coat is distinguished by the deep blackish- 
brown colour of the anterior part of the body. 

Briefly described, the colour of a typical adult male is as 
follows :— 

Dorsal aspect of anterior part of body, as far back as behind 
the shoulders, of a uniform sooty brownish-black. On separating 
the fur, two-thirds of the length of the hairs is lighter, greyish 
or dirty white near the roots, pale greyish-brown outwards, the 
outer third shading into the above-mentioned brownish-black, 
without presenting any tendency to zonal arrangement. ‘The 
posterior part of the body presents a mottled appearance, due to 
the intervention of yellowish-reddish zones. On separating the fur 
in this region, two-thirds of the length of the hair is quite uni- 
form dark; the last third is about equally divided between the 
light zone and a black terminal one. 

Singularly deep rusty-red are the rump and thighs. This is 
due to the absence of the dark terminal zone of the hairs in 
this region, each hair being black at its base, and the terminal 
third entirely rusty. Tail, arms, and feet black. 

Dorsal and ventral aspects are noticeably divided by a lateral 
rusty-red stripe of shorter hairs on each side, running from the 
arm-pit to the flank. The real median ventral stripe, wider than 
the just-mentioned lateral one, again assumes the dark brown 
colouring of the anterior part of the body. 

In the face the most salient features are the white eyebrows, 
meeting in the median line at the base of the nose, forming 
a very striking double crescent.  Circumbuccal zone whitish, 
more extensive than in all the preceding species, but not so 
sharply outlined. 

Minas PrLEAtus Is. Geoffroy et Deville. 

This species was figured in 1848 by Geoffroy in the ‘ Archives du 
Muséum,’ vol. v. pl. 31, and described under the name of “ Tamarin 
a calotte rousse,” p. 569. Up to the time of my paper read at 
Berne, at the International Zoological Congress, it seemed to me, 
judging from the literature within my reach, that it was repre- 
sented only by the single original specimen, coming from the Rio 
Javary, and preserved in the Paris Museum. (I saw it there 
some days before, without a label, stowed away on a side-shelf.) 

At that time we had obtained from the upper River Purtis two 
other specimens, a pair. This splendid species has its chief 
distinguishing marks in the cinnamon-coloured sealp-patch, the 
brownish-black general colour, and the sharply outlined white 
circumbuccal zone, including the nostrils, which are completely 
surrounded by a narrow white band. 

Being in London for the Meeting of the Fifth International 
Ornithological Congress in July 1905, Mr. Oldfield Thomas, of the 

Proc. Zoou. Soc.—1907, No. VII. 7 

98 DR. E. A. GOELDI ON MARMOSET [Jan. 15, 

department of mammals at the British Museum, to my no small 
surprise showed me a small series of skins from different points 
in the Upper Amazon region, one having been furnished by 
Dr. von Ihering, from the River Jurua expedition. 

After my return to Para I saw by the ‘ Revista do Museu 
Paulista,’ vol. vi. (1904), p. 416, which had been sent in my 
absence, that Dr. von Ihering refers to three individuals in the 
collections made by Mr. Garbe at the time of the above men- 
tioned Jurud trip. He, however, classifies it as a new subspecies 
—WM. pileatus jurwanus—which according to his statement is 
distinguished by the dark colour of the hair of the back, which in 
the description of the original type-specimen, made by Geoffroy, 
is stated to be reddish at its base. It is true, that Geoffroy expresses 
himself in the following terms :—‘“ Le corps est supérieurement re- 
couvert, ainsi qu’on l’observe si souvent chez les Hapaliens, de 
poils roux dans la plus grande partie de leur étendue, anneleés de 
blanchdtre et de noire vers la pointe. Il] resulte de cette dis- 
position, chez JZ. pileatus, un mélange de gris et de noir, mais 
non des bandes alternatives de l’une et de autre couleur.” 
Although the adjective “roux” may not be perhaps a happy 
term, the rest of the description and the figure itself indicate 
an animal which does not differ essentially from the specimens 
examined by me in London, nor from my two mounted speci- 
mens here at Para. Geoftroy evidently wrote with the desire to 
differentiate the uniform general colouring of this species (which, 
as already seen, is usually a common feature of most Amazonian 
Callitrichidee) from the distinctly striped colouring of certain 
southern Marmoset Monkeys (Hapale jacchus, &c.). 

On separating the fur of the back of my two individuals in the 
Para Museum, the hair appears to be of the same deep blackish- 
brown as in the case of WW. griseovertex. The terminal fourth 
then shows a light greyish-white zone, followed by a terminal 
nearly black one. It follows therefore that in this detail my 
Purts specimens accord with the Jurua specimens of Ihering. 
However, this detail does not seem to me of sufficient weight to 
justify the establishing of a separate subspecies. 

MipAs MyYsTAX Spix. 

Acquaintance and description of this species date from the 
same period and come from the same source. Spix brought his 
specimens from the River Solimoes; a female is figured on plate 22 
of his work. Our Museum possesses a considerably darker male 
specimen from the River Jurua. 

Instead of entering into a detailed description, I would simply 
call attention to the characteristic fact, that the white circum- 
buccal zone is nearly identical im extent and form with that of 
the preceding Midas pileatus, but with a stronger tendency to 
form a moustache. 


Mipas 1LLicert Pucheran. 

I have recently received, through the kindness of the wife of 
the German Consul at Para, Dr. Olshausen, an example of another 
species of Amazonian Marmoset Monkey hitherto unrepresented 
in our collections. It had been obtained at a very early age from 
an Indian woman at Iquitos, and kept as a pet until its mistress 
was about to sail for Europe. It lived several months at the 
Museum until about half-grown, and is now mounted. 

Dorsau Aspect.—Principal colour a lovely dark brown-red 
covering the nape of neck, shoulders, and outer side of arms and legs, 
embracing with a small band the rump and base of the tail. The 
back, however, properly speaking, forms an exception to this colour, 
bearing a very long oval patch, distinctly outlined, of dirty black 
in the centre and greyish borders, with the tendency to form 
posteriorly dimly apparent light and dark transverse bands. Head, 
hands, feet, and tail black. 

VENTRAL ASPECT.— Uniformly brownish-black from the throat 
as far as the anus, embracing the inner side of arms and legs, 
separated from the dorsal patch by a brownish-red margin. 

In the black face we again find the white circumbuccal zone. 
But in this case it leaves the nostrils free and the white runs up 
to the cheeks in a triangular form nearly as far as the outer 
corner of the eye. 

By this feature the animal immediately proves to belong to 
the group 6, subdivision a of the classification of Schlegel (‘ Singes,’ 
p. 262), that isto say tothe Hapale devillei group. Among all the 
figures I have at my disposal, my specimen corresponds best with 
the animal represented in plate 13 of the ‘ Proceedings of the 
Zoological Society,’ 1871, by Bartlett, with the designation of 
Midas devillei $ (considerably better than with the figure of 
M. devillei, fig. 3, plate 6, in the ‘ Atlas’ of Castelnau, which lacks 
any trace of a distinctly coloured dorsal patch). As the animal 
there represented is attributed by Schlegel (‘ Singes,’ p. 263) and 
by Forbes (‘Handbook of Primates, p. 145 seq.) to Hapale 
illigert Pucheran (‘ Revue de Zoologie, 1845, p. 336), and as the 
description of this species given by these two authors coincides 
satisfactorily with my Iquitos individual, I think I have to do 
with a half-grown specimen of Midas illigeri Puch.,a Marmoset 
Monkey stated to be fairly abundant in the Peruvian Amazons. 



February 5, 1907. 

His Grace Tue Duke or BeprorD, K.G., President, 
in the Chair. 

Mr. F. Martin Duncan, by permission of the Charles Urban 
Trading Co., Ltd., gave a cinematograph exhibition of animals in 
the Society’s Gardens and other zoological subjects, chiefly the 
life-history of Insects. 

Mr. Oldfield Thomas, F.R.S., F.Z.S., exhibited a collection of 
Mammals and Birds from the Islands of Saghalien and Hokkaido, 
N. Japan, made by Mr. Malcolm P. Anderson in carrying out the 
Duke of Bedford’s Exploration of Eastern Asia. Mr. Thomas 
proposed to give a full account of the Mammals on a later 

Dr. W. T. Calman, F.Z.S., read a paper entitled ‘On new or 
Rare Crustacea of the Order Cumacea from the Collection of the 
Copenhagen Museum. Part I. The families Bodotriide, Vaun- 
tompsoniide, and Leuconide.” b 

This paper will be published entire in the ‘ Transactions.’ 

The following papers were read :— 

1. The Origin of the Lateral Horns of the Giraffe in Foetal 
Life on the Area of the Parietal Bones. By H. Ray 
lankestprs) WIA. DiSe., use, SEeRES* HeZzas:, 
Director of the Natural History Departments of the 
British Museum. 

[Received February 5, 1907. ] 
(Text-figures 24-36.) 

A remarkable and wide difference between the Giraffe and the 
Okepi is constituted by the position and relation of the lateral 
horns in these two animals in regard to the bones of the skull. 
AsI pointed out in my memoir on the Okapi read in 1901 (Trans. 
Zool. Soc. vol. xvi. p. 279), the bony horn-cone of that animal is 
attached to the frontal bone, and it is the frontal bone which is 
raised into a boss for its support, whilst even in the hornless skulls 
supposed to be those of the female these frontal bosses are present. 
On the other hand, in the young Giraffe the main axis of the 
lateral “ossicone”* falls within the area of the parietal bone 

* T use the term “‘ ossicone ” in the present paper for the independently ossifying 
bony cones which are found in Okapi and Giraffe on the frontal and parietal areas 
and in the Giraffe also in a median position. In my memoir of 1901 I spoke of such 
structures as “ossicusps,” a term which I now wish to apply more generally, 

reserving the term “ossicone” for the peculiar separately ossifying cones of the 

1907. ] HORNS OF THE GIRAFFE. 101 

(text-fig. 24). The wide-spreading base of the cone-like ossicusp 
subsequently encroaches, it is true, over a large portion of the 
frontal bone. In the adult both the parietal and the frontal are 

Text-fig. 24. 

Lateral view of the skull and lower jaw of a very young Giraffe, measuring 
30°8 centimetres from the occiput to the anterior border of the premaxilla: 
preserved in the British Museum. The drawing is five-twelfths of the natural 

oce., occipital crest; gé./., Giraffine conical tumescence of the parietal bone, above 
which is developed the lateral ossicone (p.oss.); sfp., the fronto-parietal suture; 
ot.l., position of the lateral tumescence of the Okapi, ‘absent here ; gt.m., position 
of the median frontal tumescence of the Giraffe, which in this young specimen 
is still entirely undeveloped ; o¢.m., position of the median tumescence of the 
Okapi’s skull (basinasal) ; pln, prelacrymal vacuity; can., bifoliate canine 
(deciduous dentition). 

(From Trans. Zool. Soc. vol. xvi. p. 293.) 

enlarged and tumescent, and both enter largely into the com- 
position of the lateral horn of the adult Giraffe (text-figs. 25 & 33). 
The whole form of the skull is rendered different in the two 
genera by this relationship of the ossicone to the frontal exclusively 
in the one, to the parietal primarily but not exclusively in the 
other (see text-figs. 26 & 27). 

In a skull of a very young Giraffe (text-fig. 24), probably about 
a year old, preserved in the British Museum, the lateral ossicone 
is seen to rest almost entirely on the parietal. A transverse 
section (text-fig. 28, p. 104) shows that the anterior margin of the 
enlarging base of the bone constituting the ossicone has spread— 


Text-fig. 25. 

Lateral view of the skull of a Giraffe, about two-thirds grown. 

oce., occipital crest ; p.oss., parietal ossicone (epiphysis) overlying the parietal conical 
upgrowth and spreading on to the frontal bone; sfp., fronto-parietal suture ; 
o.t.l., position of the lateral frontal tumescence in the Okapi, absent here ; 
g.t.m., the characteristic median tumescence of the Giratte’s frontal, devoid in 
this specimen of any secondary cap or epiphysis, absent in the Okapi; pl.v., the 
prelacrymal vacuity. 

(From Trans. Zool. Soc. vol. xvi. p. 284.) 

Text-fig. 26. 

(yyyt 4 

= ihe Le 
) vt iM) GH, 
in tat 

View from above of the fronto-parietal region of the skull of a very young Giraffe. 
The parietal epiphyses were already ossified, but separable, and are here removed. 

occ.h., exostosis of the occipital crest, which in some adult Giraffes forms a second 
pair of “horns” (five-horned Giraffe) ; gtl., the Giraffine lateral tumescence, seen 
here to originate in the parietal bone, which carries the conical ossicone aud forms 
the Giraffe’s paired “horns ”’; o¢./., the position of the Okapi’s lateral tumes- 
cence of the frontal bone, absent in Giraffe; zy., zygomatic arch; po., posterior 
angle of the orbit ; ao., anterior angle of the orbit; pl.v., prelacrymal vacuity 
sof., supraorbital fossa; sfp., fronto-parietal suture. 

(From Trans. Zool. Soe. vol. xvi. p. 291.) 

1907. | HORNS OF THE GIRAFFE. 103 

invaded as it were—the area of the frontal bone to a very slight 

It seemed hardly possible to doubt that the ossicone of the 
Giraffe takes its origin within the area of the parietal bone, but 
that conclusion was forbidden by the explicit statement of the late 
Sir Richard Owen who, in a paper published sixty-seven years ago 
(1840) in the ‘ Transactions’ of the Zoological Society, described a 
newly-born Girafte which had died in the Gardens of the Zoological 
Society. Owen there states that he found the lateral horns of this 
Giraffe to be definitely attached to the frontal bone, and to that 

Text-fig. 27. 


View from above of the fronto-parietal region of the skull of an immature Okapi. 

oce.h., angle of the occipital crest ; g.¢.J., position of the lateral tumescence of the 
parietal which supports the paired ossicones of the Giraffe, absent here; 0.t.l., 
the Okapian tumescence of the frontal which supports the paired ossicones of 
the Okapi; zy., the zygomatic arch; po., posterior angle of the orbit; ao., 
anterior angle of the orbit ; pl.v., prelacrymal vacuity; ofm., the slight median 
tumescence of the base of the nasals of the Okapi; sof., supraorbital fossa; 
sfp., the fronto-parietal suture. 

For comparison with text-ficure 26. 
(From Trans. Zool. Soe. vol. xvi. p. 290.) 

bone exclusively. He gives the drawing, which is copied in text- 
fig. 29, p. 104. He draws attention to the suture (s) separating 
the two bones seen in section, and he states that # is the frontal 
boneand y the parietal. He arrives at the conclusion that whilst 
the lateral horns of the Giraffe are seen thus to originate as do the 
horns of all other Pecora, in connection with the frontal bone, 


yet that in the Giraffe the growing horn must spread from its 
original position, and in fact take up a new position on the 
parietal, with which he recognises that it is largely in contact 
in adult life. 

Text-fig. 28. 

é N 

y 6 \ 

i 2 RS 
if ‘ LN 

Bes ESN 

ea oo" 7). 088< 

Text-fig. 28.—Sagittal section through the bony tissue of the ossicone and the roof 
of the skull of a very young Giraffe (same specimen as that drawn in text- 
fig. 24). Drawn of the natural size. 

p.oss., the ossicone; s., the parieto-frontal suture; a:., the parietal bone; 
y., the frontal bone. 

Text-fig. 29.—Copy of the drawing (natural size) of a sagittal section through the 
ossicone and the roof of the skull of a newly-born Giraffe, published by_the late 
Sir Richard Owen in the Trans. Zool. Soc. 1840. 

p.oss., the ossicone; s., the parieto-frontal suture; «., stated to be a “ frontal” by 
Owen but actually parietal; y., stated to be “ parietal”? by Owen but now shown 
to be frontal. 

_ I felt considerable doubts as to the correctness of this observa- 
tion, and obtained through the kindness of Prof. Stewart some 


five years ago the actual skull of the newly-born Giraffe examined 
by Owen, and still preserved at the Royal College of Surgeons. 
The whole of the frontal and parietal regions had been cut away 
from the skull and the pieces could not be found. There was 
no evidence to be obtained from the specimen as to what really 
was the nature of the bone a (in text-fig. 29) and the bone y. I 
formed the hypothesis that Owen had had the horn-bearing region 
cut out and a section made by an assistant. The section is that 
which he figured and is here copied. But the piece having been 
detached from the rest of the skull, Owen seems to have mistaken 
right for left and back for front, so that in reality the bone 
marked a is the parietal and the bone marked y is the frontal ; 
and the young horn or ossicone is resting on the parietal as it 
does in the later stages of growth, and not on the frontal as 
supposed by Owen. 

I could not test the truth of this hypothesis without examining 
myself a newly-born Giraffe or a well-advanced fcetus, and accord- 
ingly I have made efforts to obtain such a specimen by application 
to the officials of the late African Department of the Foreign 
Office and to those of the Colonial Office, as well as to naturalists 
and sportsmen. Nonewly-born or feetal Giraffe came to hand, nor 
could I hear of one as being preserved in any Museum in Hurope. 
Accordingly I was very grateful when last summer our Secretary 
was able to place at my disposal the fcetal Giraffe which was 
removed from its mother after her death in the Gardens in April 
1906. This feetal Giraffe was figured and described in a general 
way by Mr. Beddard (Proce. Zool. Soc. 1906, p. 626), but the 
examination of the skull was kindly left to me. The dead 
mother of this foetus was a South-African (Transvaal) Giraffe 
(G. camelopardalis wardv) and the father a Kordofan specimen 
(G. camelopardalis antiquorum). Mr. Beddard has given the 
dimensions of the foetus, and has estimated that it had probably 
completed two-thirds of its fcetal life. He has pointed out and 
figured the large size of the incipient lateral horns, and their 
free extremities tufted with long hair, and has noted that their 
substance 1s of a gristle-like consistency. 

Soon after I received the specimen, the integument (C) was 
reflected from the right side of the fronto-parietal region of the 
head, under my supervision, by my assistant Dr. Ridewood ; and 
by subsequent reflection of part of the periosteum the view 
obtained which is given in text-figure 30. The integument of 
the right side of the head (C) was thrown back, and the periosteum 
of the parietal bone was reflected (D) excepting that part lying 
beneath and forming the base of the right lateral horn. ‘This was 
pinned down and cut away from the rest of the periosteum, leaving 
it as an oval area A, marking exactly the position of the ossicone 
(fibrous and soft) on the parietal bone. In the drawing the suture 
separating the frontal from the parietal bone is seen (szé.), and it 
is demonstrated that the base of the young lateral horn or ossicone 
is wholly within the area of the parietal bone, to the periosteum 
of which it is loosely attached by connective tissue. 

[Feb. 5, 



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‘OE “SU-9x9], 

1907. ] HORNS OF THE GIRAFFE. 107 

The substance of the young ossicone was free from osseous 
deposit but of a tough consistence. Microscopic sections showed 
it to consist of a fibro-trabecular tissue with abundant interspersed 
cellular elements. 

The dissection as here presented definitely establishes the fact 
that the site of origin of the lateral horns or ossicones of the 
Giraffe is entirely within the area of the parietal bone. The 
supposition that Sir Richard Owen had by inadvertence reversed 
the cut-out portion of the fronto-parietal area of his newly-born 
Giraffe, and had thus identified parietal as frontal and frontal as 
parietal, is confirmed. The importance of the distinction between 
the genus Giraffa and the genus Okapia, arising from the parietal 
position of the lateral ossicones in the former and their frontal 
position in the latter, is thus placed on a firm basis, since it 1s 
shown that at a stage of growth even earlier than that of birth 
the “forecast” or rudiment of the Giraffe’s lateral bony cone 
(ossicone) is placed and attached absolutely and solely within the 
area of the parietal bone. 

Text-fig. 31. 

* i l) \ 

LZ i [Yf ye 

The left “ horn ” of the fcetal Giraffe, drawn of the natural size. 

A. Seen from behind (postero-external face). 
B. Seen from in front (antero-internal face). 

The form of the soft unossified forecasts of the lateral ossi- 
cones in this foetal Giraffe is worthy of further notice. As shown 
in Plate V. accompanying a later paper in this volume, and in the 
text-figure here given (fig. 31), the upgrowth is of considerable size, 
is compressed so as to give a narrow oblong area in transverse 
section, and is set on the head so that the elongated basal area 
has an oblique position, with its long diameter directed back- 
wards and inwards towards the median line. The surface of the 
ossicone-forecast is covered with hair, which 1s very coarse and 
long at the free upstanding margin, as shown in text-fig. 31. 


The text-figure 32 also shows that three of the dark hair-bands 
are disposed around the base of the young ossicone. The great 



Right Horn 



Diagram to show tbe flattened plate-like form and the orientation of 
the horns of the foetal Giraffe. 

a, b, c, d. The four left-side inter-cornual colour-bands of the pelage. 

lateral compression or flattening of the young structure is 
remarkable as compared with the fuller circular contour of a 
transverse section after ossification has advanced. 

Possible Relations of the Giraffid Ossicones to the bony Horn- 
cores of Bovide and the Antlers of Cervide. 

In my memoir describing Sir Harry Johnston’s specimens of 
Okapi, I ventured on some speculations as to the relationships of 
the bony growths called horns and antlers in the Pecora. These 
speculations were vitiated by the uncertainty of existing know- 
ledge as to the actual embryological origin of the structures 
compared. They assumed the origin of the horn-cores of Bovidee 
as separate rudiments which become united to the osteogenetic 
tissue of the frontal bone at an early period of development. 

Although convincing histological accounts of their development 
are not yet in existence, it seems to result from the observations 
of Dirst that the horn-cores of Bovide are not of independent 
origin, but are actual outgrowths of the osteogenetic tissue of the 
frontal bone. The same origin appears to hold for the horn-style 
or column which supports the antler of the Cervide. 

It is unfortunately the fact that our knowledge of the early 
embryological history of the ‘ossicones” of the Giraffe and 
Okapi is still more defective. All we know is (1) that in Owen’s 
newly-born Giraffe the rudimentary unossified ossicone was sepa- 
rated (as shown in text-fig. 29) by the dense periosteal membrane 
from the subjacent parietal bone; (2) that in the feetal specimen 
of two-thirds time here described the soft forecast of the ossicone 
was equally cut off from the subjacent parietal bone by dense 

1907. ] HORNS OF THE GIRAFFE. 109 

periosteum ; and (3) that in a much younger dried skull there is 
no indication on the cranial wall of any “ outgrowth.” 

We shall not be ina position to speak definitely until other 
foetal Giraffes of younger stages have been examined by proper 
histological methods; but it seems legitimate to consider the 
lateral ossicones of the Giraffe, and therefore of the Okapi, as 
originating in a fibrous osteogenetic mass which gives rise to a 
protrusion of the integument and originates in the connective 
tissue of the integument rather than in the osteogenetic tissue of 
the cranial roof, from which it is separated by a dense membranous 

The lateral ossicones of the Giraffe appear to have the same 
nature and mode of origin as has the centrally-placed ossicone of 
the same animal. This median ossicone is variable in size, and 
does not appear until many years after birth when growth is 
nearly complete. The histological processes by which the median 
ossicone is formed have never yet been studied, but it is practically 
certain that it forms not as an outgrowth of the bone of the 
cranial roof, but as a “dermal” or tegumentary growth external 
to and independent of the cranial bone. 

The same process which leads in the Giraffe to the formation of 
a median ossicone, In some cases becomes specially active and leads 
to the formation of additional “ supernumerary ” sub-tegumental 
ossifications. Thus two such of smaller size than the normal 
median ossicone are seen in the median line in a skull of Giraffe 
in the British Museum, in which bony deposits on the margin 
of the orbit are also seen. The skull of Okapi brought by 
Capt. Boyd Alexander from the Welle River shows such an 
exceptional sub-tegumental bony deposit on the margin of each 
orbit ; and it is to be noticed that both in Giraffes and Okapis in 
old individuals the base of each ossicone spreads very widely as a 
thin encrusting layer, so as to involve much of the frontal in 
specimens of Giraffe and a large surface of the parietal in the case 
of Okapi. 

These superficial ossicones and their outspreading marginal 
growths of the Giraftide cannot be accurately marked off in later 
life, although they can be separated in earlier life, from the highly 
important upgrowths or “tumescences” (as I previously called 
them) of the cranial bones over which they lie. The completed 
lateral horn of the Giraffe consists very largely of a conical up- 
growth of the parietal bone, and also of the frontal bone—occupied 
by an air-sinus—developed between the tabule of the bone. The 
basal region of the upgrowth involves the frontal bone more 
largely than the parietal. The originally independent ossicone 
contributes but a small amount to the whole bulk of the structure. 
It forms merely the terminal knob, and is fitted over the tumes- 
cence like a superficial investment which dwindles in thickness as 
it descends the cone until it becomes a mere film. 

The position of the lateral and median ossicones in an adult 


(though not senile) Giraffe, and the relation of the sinus of the 
parietal and frontal to the lateral horn, is shown in the figures (text- 
figs. 33 & 34) which were prepared from photographs of sections 
of a skull made under my direction. It will be observed that it 
is not possible in such sections to distinguish the line of ankylosis 
of the separate bones; we can only guess somewhat vaguely as to 
what belongs to each of the three elements fused together, viz., 
ossicone, parietal, and frontal. 

Text-fig. 33. 

Skull of Giraffe, left side; key-figure, a little larger than one-sixth (linear) of the 
natural size, showing the directions in which the skull was cut. 

(From the Proc. Zool. Soc. London, 1904, vol. i. p. 151.) 

Thus we see that the “‘ossicone” of the adult Giraffe is essen- 
tially but a cap of bony substance fitting over the great upgrowth 
or conical tumescence of the cranial wall—at the first most marked 
in the parietal area,—and that it makes its first appearance as a 
solid growth of fibrous tissue resting on the flat cranial roof. 
The same conclusion may legitimately be drawn from what we 
can see of the early and later conditions of the ossicone in 


* The tumescence of the frontal bone of each side in Okapi, which ankyloses with 
the ossicone overlying it, is of considerable volume, like those to which the parietal 
gives rise in the Giraffe. I shall have an opportunity of describing this structure 
more fully hereafter. At the present moment I desire to draw attention to a curious 
fact with regard to the rudiment of a median horn in Okapi. The base of the nasal 
bone gives rise to a small but well-marked median tumescence in the Okapi. I 
described this in Sir Harry Johnston’s specimen (Trans. Zool. Soc. vol. xvi.). In horn- 
bearing skulls of Okapi this is more pronounced than in the hornless speeimens ; it 


Since it is clear then that the Giraffe’s horns comprise two 
bony factors, the question arises which of the two corresponds 
with the horn-core of Bovide, or whether either or neither had 

such a correspondence. 

Text-fig. 34. 

we3)s, parrect 
\ horn 

2 ul SGM 


alisph ; “y Vi ‘ 

7 | 
bus sph .  pteryg. 

View, looking forwards, of the skull cut in the direction of the line 4 in text-fig. 33. 
One-third (linear) of the natural size. 

(From the Proc. Zool. Soc. London, 1904, vol. i. p. 154.) 

is more strongly marked in the horn-bearing skull of Okapi brought by Capt. Boyd 
Alexander from the Welle River than in any other specimen seen by me. No “ossi- 
cone” or tegumentary cap has been observed in Okapi in connection with this 
median tumescence. Has there been one present in the ancestors of Okapi? Or 
does the tumescence precede the formation of a tegumentary ossicone ? 


Text-fig. 35. 

i] te £ 

Bi) 4 Oi 
Wy; ip Chis ae 
MM 27 

KOEN 7 / 

Drawing, of the natural size, of a young foetal skull of a. Giraffe, preserved in 
the British Museum. ‘he foetus is at so early a stage that no trace of the 
future horn or ossicone is presented, and moreover the arietal bone does not 
form the vertex of the brain-case as in the skull drawn in text-fig. 24, p. 101. It is 
worthy of remark that a line drawn from the occ.-par. suture to the fronto-par. 
suture is in this early foetal skull almost at right angles to a line drawn from 
the latter to the naso-frontal suture, a condition which is even more strikingly 
exhibited in the adult skull of the Common Ox (Bos) as contrasted with that of 
Ovis, Ovibos, and Antilopide. In adult Giraffe the surfaces of the frontal and 
parietal are more nearly parallel, and in Okapia quite so. It is a very curious 
fact that in Bos the whole posterior region of the brain-case formed by the 
parietals, which in Okapia stretches horizontally backwards from the fronto- 
parietal suture to a distance as great as that occupied by the frontal portion, is 
abolished! The parietals are vertical and not horizontal, and a sort of false 
occipital ridge is formed by the fronto-parietal suture. The very young Giraffe- 
foetus approaches this condition. 

Tn answer to this question, it appears to me that the following 

statements are justified :— 

1. Since the bony horn-core of Bovide originates as a part of 
the osteogenetic tissue of the frontal bone, it cannot 
(according to our present knowledge) be identified without 
considerable qualification with the free tegumentary t 
ossicones of Giraffide. 

2. The upgrowth (tumescence) of the frontal bone in Okapi and 
of the parietal and frontal in Giraffe, which forms the 
bulk of the lateral horns in those animals, is more nearly 
similar in nature to the bony horn-cores of Bovide than 
are the ossicones of Girattidee. 

3. The frontal tumescence or upgrowth of Okapi cannot be 
considered as morphologically identical with the parietal 
tumescence or upgrowth of Giraffa. 

4. The free lateral ossicone of Giraffa might legitimately be con- 
sidered as morphologically identical with the free lateral 
ossicone of Okapi. The free tegumentary ossicone of an 

1907. ] HORNS OF THE GIRAFFE. 113 

ancestral form might in a series of generations shift its 
position from the frontal to the parietal area ; and it might 
reasonably be admitted that the upgrowth or tumescence 
of the frontal ceased to develop when the parietal position 
was assumed by the ossicone with consequent tumescence 
and upgrowth of the parietal bone. 

5. On the other hand, the theoretical assumption that the 
frontally-placed ossicone of Okapi and the parietally-placed 
ossicone of Giraffe are independent of one another and 
possibly co-exist in an ancestral form, is favoured by the 
fact that the Giraffe does develop a third well-marked 
ossicone in the mid-line of the frontal, and that both 
Okapi and Giraffe exhibit minute supernumerary growths 
of the kind on the cranial surface. 

6. It results from these considerations that it is not possible at 
present to trace the lateral horns of the Okapi and the 
Giraffe into any close genetic relationship with those of 
Bovidee—still less of Cervide. At the same time it is 
possible that the peculiar superficial element of the bony 
horn (the ossicone) is identical in the lateral horns of 
Okapi and Giraffa, having shifted its position backwards 
in the latter genus. This conclusion is not, however, by 
any means forced upon us since the Giraffide are known 
to have an additional ossicone—the median one; and it is 
therefore not without analogy that independent frontal 
and parietal G sicones should develop. 

I am aware that it is not difficult to make assumptions by 
means of which a genetic relationship between the lateral horns 
of Giraffide, Bovide, and Cervide is rendered possible; but it 
should, I think, be clearly understood that there is at present no 
direct evidence to support these assumptions. It may be assumed 
(a) that a bony horn of the nature of the horn-core of the Bovide, 
or of the antler-column of Cervide, has in some remote ancestors 
of the Giraffidee become segregated from the frontal bone of which 
it was a part, and acquired independent existence as a fibrous 
rudiment as well as independent ossification, thus establishing the 
independent lateral ossicone of the Giraffide. Or, again, it may 
be assumed (6) that in ancestors of the Bovide and Cervide, 
bony horns which were existing as free tegumentary products, 
ankylosing in mature age with subjacent cranial bones, became 
so ankylosed at earlier and earlier stages of development until all 
trace of their independent origin was lost, and they appeared to 
originate as growths of the frontal bone itself. The stock so 
endowed gave rise (it would be assumed) to Bovide and Cervide ; 
that portion of the ancestry which retained the original method 
of development of free tegumentary ossicones became, on the other 
hand, the progenitors of the Giraftide. 

I am not aware of any facts in the structure of living or extinct 
Artiodactyla which furnish an analogy for either of these pro- 
cesses of transformation. Nor do I think that our knowledge of the 
extinct forms such as Samotheriuwm, Helladotherium, Sivatherium, 

Proc. Zoou. Soc.—1907, No. VIII. 8 


Bramatherium, and Hydaspitherium, is of a nature to assist in 
establishing the existence of such a remarkable transformation. 

The assumption a—namely, that the transition was from frontal 
outgrowths such as Bovide and Cervide present, to the segregation 
and independence of the Giraffid ossicone—meets (it may be 
pointed out) with an almost insuperable obstacle in the free median 
frontal ossicone of the Giraffe, for which there is no forerunner 
in the Bovi-Cervine scheme of solid continuous outgrowths of 
the eranial wall. The consideration of the Lower Miocene Proto- 
ceros with its four pairs of horns does not help us much in this 
question, though its possession of a pair of parietal and of two 
pairs of frontal upgrowths or ‘ bosses’ is significant. 

Various Positions occupied by the Paired Frontal Horns of 
Cavicorn and Cervine Ruminants. 

An interesting fact in relation to the question of the identifi- 
cation of Giraffe’s parietal with Okapi’s frontal ossicones is that 

Text-fig. 36. 


Anterior horns 

oy Be of Tetraceros. 

= a 
Antilocapra americana ~ 
Qreotragus saltator. 

__Ovibos moschatus, 
Cervus elaphus. 

Ovis tragelaphus __ _ _ 

Nemorrhedus goral. _ Posterior horns 

of Tetraceros. 

“Alcelaphus caama. 
Cephalophus coronatus. 


Diagram to show the various positions on the frontal bone at which the bony horn- 
cores of the Cavicorn and Cervine ruminants may take their growth. Though 
ranging widely over the frontals these upgrowths are never found as part of the 
parietal bones. 





( Half the natural size.) 


the paired frontal horns of Antelopes, Bovines, and Deer occupy 
very widely-separated positions in different genera. Dr. Ridewood 
has kindly prepared for me a diagram (text-fig. 36) showing these 
varied positions in a series of genera. The most remarkable 
position is that of the horn-cores of the Antelope Cephalophus, 
which. is not fully exhibited in the diagram. The frontal in 
some species of this genus actually pushes out a process into the 
area of the parietal, upon which the horn-core rests. The horn- 
cores are seen (by reference to the diagram) to arise sometimes at 
the hinder margin of the frontal, sometimes on the orbital ring, 
sometimes in the anterior third of the frontal. The co-existence 
of two pairs in Tetraceros suggests a multiplicity of horns in 
ancestral forms. 

The position of the horn-cores in the genus Bos is not marked 
in the diagram ; it is identical with that of the posterior pair of 

2. Parallel Hair-fringes and Colour-striping on the Face of 
Foetal and Adult Giraffes. By HE. Ray LANKEsTER, 
VeAn Osc. tO. RIS. EZ.S., Direction ot the 
British Museum (Natural History). 

[Received February 5, 1907. ] 

(Plate V.* and Text-figures 37-48.) 

When examining the feetal Giraffe which I received from the 
Society in the past summer, I observed a number of parallel 
bands or stripes of dark and light colour on the hairy coat of the 

Text-fig. 37. 

Left side of the head of the foetal Giraffe described, showing colour-stripes on the 
snout and above and below the eye. The small arrows indicate the direction of 
slope of the hairs. About three-tenths of the natural size. 

* For explanation of the Plate, see p, 125. 



face—between the two lateral horn-sacs, also between the eyes and 
above the eyes, which are carefully represented in the coloured 
drawing (Plate V.) of the face of the foetal Giraffe. I observed 
sinilar but more strongly marked and broader bands of alternating 
reddish-brown and paler colour between the nostrils (see Plate V. 
and also text-fig. 30, swpra, p. 106) and at the side of the upper 
lip, on the front of the lower jaw and below the eye (text-fig. 37, 
p. 115). 

rae colour-bands between the horns and the eyes varied in 
intensity according to the angle of the incident light, and could 
be temporarily destroyed by pressing the skin and smoothing 
down the hair. After careful examination of the hairs, I came 
to the conclusion that there was no actual difference of colour in 
the hairs occupying the darker stripes and those placed on the 
lighter tracts, but that the phenomenon was due to the existence 
of parallel linear depressions or wrinkles the existence of which 
was made obvious by a transverse section of the integument 
(text-fig. 38). The hairs are crowded together in the trough of 
the wrinkle, and further apart in the convex intermediate areas. 
That “ wrinkling” could produce such an impression of dark and 
light banding was demonstrated by the casual folds and wrinkles 
of the integument on the legs, and by purposely producing such 
wrinkles by pressing or folding the hairy integument. 

Text-fig. 38. 

Sister a = = 
Su oy a eee 2 
yp = — Ge = = 
kee (ASS ae eS SS ea 
ws Do ES SE 8p SS SS Se 
ee NS SE Se == 
= SSS 

Section across three dark-coloured bands above the eye of the foetal Giraffe, showing 
three longitudinal furrows or depressions in transverse section corresponding to 
the dark bands. Magnified. 

Nevertheless I could not attribute the colour-banding to a 
mere accidental or casual formation of wrinkles. Their definite 
form and arrangement precludes such an explanation. They 
appeared to me to be the expression of a definite structural 
condition. Moreover, immediately over the eyes and on the 
snout the difference of colour of the alternating bands was very 
strong, the darkly-coloured stripes being of a strong reddish- 
brown tint and the intermediate bands quite pale; and in this 
case a pigment was present in the hairs of the darker stripes 
which was not so richly developed in those of the lighter 
neighbouring stripes. I found, on microscopic examination of 
the hairs, that they could be roughly divided into three sizes ; 


the largest, few in number ; medium-sized, more numerous; and 
a smallest size, the most abundant (text-fig. 39). The two larger 
sizes of hair were remarkable for appearing almost colourless in 
their lower (proximal) moiety and very dark in the upper (distal) 

Text-fig. 39. 

[Be iB 7 

Section of the skin of the frontal region of the foetal Giraffe, showing 
three sizes of hairs. Greatly magnified. 

moiety. The smallest hairs were dark-coloured throughout. 
When a piece of the hairy colour-banded region was examined 
with a strong lens, the tips of the hairs were seen to converge from 
the two sides of the shallow furrows or wrinkles as shown in the 


text-figure 40. The superposition of the darker free ends of 
the hairs conceals theix paler basal regions, and thus intensifies 
the difference between the apparent colouring of the troughs or 
wrinkles and the intermediate spaces where the hairs do not 
converge. An interesting experiment in regard to this matter was 
made by my assistant Dr. Ridewood. He took a piece of pale 
hat-maker’s plush, and stained with dark pigment the free ends 
of its hairy surface. He found that on throwing this manufac- 
tured material into a series of wrinkles, very strong alternation 

Text-fig. 40. 


SS — 
—— SS 



Surface-view of banded “pelage”’ of the frontal region of foetal Giraffe. showing the 
convergence of the hairs at the three longitudinal bands of dark appearance. 

of dark and light colour-stripes could be produced. (This pre- 
pared material was shown to the Meeting and dark and light 
bands produced in it and removed at pleasure by alternately 
throwing it into wrinkles and stretching it so as to remove the 

Although these colour-stripes on the head of the fcetal Giraffe 
thus appear not to be due to alternate tracts of hair of differing 
colour, it seems to me that they have a real existence as effective 
colour-marking, and that their structural cause is to be found in 
the differentiation of the attachment of the panniculus carnosus 


Text-fig, 41. 

An enlarged drawing of a single hair of average size from the fr 

ontal region of an 

adult Giraffe (Giraffa camelopardalis wardi), showing the dark pigmented 

free extremity and the opaque white lower portion. 


(ih a 
_— a 1 
aah Za 

= £m 
Za Zi 

= = F 

Ny i i iN 

Colour-striping on the face of an adult male Giraffe from Kordofan (G. c. antiquorum). 
rawn from a specimen living in the Society’s Gardens. 

Wes Text-fig. 43. 


A nN *& 
TT i: 
BZ o 
B oI 
ZB i " 

a co | 

th Ay NG & 


Colour-striping on the face of an adult female Giraffe from Kordofan (Gc. antiquorwm). 
rawn trom a specimen living in the Society’s Gardens. 


to the integument, and in the related direction and convergence 
of the hairs in definite bands or hair streams. An examination 
of the adult Giraffe establishes the truth of this view. I visited 
the Giraffe-house at the Society’s Gardens in order to examine 
the father of the feetal specimen in which I had observed this 
colour-striping. The father is a Kordofan specimen, and there 
is also in the Giraffe-house a female Kordofan Giraffe. I was 
not a little surprised to find very strongly-marked colour-striping 
over the eyes of both these Giraffes, especially well-marked in 
the female. Portions of the muzzle are also banded, and between 
the eye and the angle of the mouth are developed from four to six 

Text-fig. 44. 



ie & | 
Set il 
Bat ae 






Colour-striping on the face of Giraffa camelopardalis cottoni from Mt. Elgon, 
Uganda. This specimen shows a horn-like exostosis over the right eye. Drawn 
from a specimen in the British Museum. 

strongly-marked horizontal colour-bands which are not present in 
the foetus (see text-figs. 42 & 43). Ifound that when the eye was 
shut and the skin above the eye stretched, the strongly-marked 
dark and light bands disappeared giving place to an irregularly 
blotched appearance *, which immediately resolved itself into 
alternate dark and light bands when the eye was opened and the 

* It seemed to me that there were in this region definite tracts in which the 
pigmentation of the hair was pale, and others m which it was dark, but exact 
observation was, I found, impossible in the living animal. 


superciliary region thrown into the normal condition assumed 
when the animal is alert. I also noticed that though the 
horizontal bands between eye and mouth never actually disappear, 
they are intensified by a muscular contraction resembling a 
sneer which sometimes is exhibited by the Giraffe. I found that 
the third Giraffe in the Gardens (a West African specimen) did 
not exhibit any colour-bands on the face. 

Text-fig. 45. 

Colour-striping of the face and muzzle of a Transvaal Giraffe (Giraffa camelopar- 
dalis wardi). Drawn from a specimen in the British Museum. 

I next proceeded to examine from this point of view the 
fine series of Giraffe heads and necks exhibited in the public, 
gallery of the British Museum, as well as several unmounted 
skins. I found that several of these specimens exhibit colour- 
banding over the eye and some of them below it, whilst these same 
specimens exhibit strong horizontal “fringes” or banding of the 
hair between the eye and muzzle, and some also show banding on 
the lower lip (see text-figs. 44-48). On the other hand, some of 
the specimens exhibit little or no trace of these bands. In none 
have I found any trace of the bands in the mediad position 
between the horns and between the eyes, shown by the feetal 
Specimen drawn in Plate V., which itself shows no trace of the 
pre-orbital horizontal stripes. 


The bands of alternate light and dark colour are as much 
larger and wider in the adult Giraffe as are all its dimensions 
larger than those of the fetus. But a fact in regard to the 
banded appearance of the hairy coat of the face has come to light 
in the case of the adult skins which is indicated in the enlarged 
drawing of a piece of the foetal pelage (text-fig. 40, p. 118). This 
is that the bands are essentially due (at any rate, those which are 
peculiar to the adult and most strongly-marked, viz., the hori- 
zontal pre-orbital group of bands) to a differentiation of the hair 
into parallel tracts of more Coase placed hairs, the points of 

Text-fig. 46. 

Wy, co 
AY WHI HIN\\, \\ WAY Ay 

Ml ea 

TAK Ain 
yt UG We 
(eee wt UY, 


nN SS 
h SW 
cc AS iN \ ND Wy 

att vif) Wy 
\) \ Wie Wy 

if Mil 

ee Hf Uf 
if} f Uji 
a Huh thi yh Hy 

The same specimen as that shown in soars 45. in this drawing the colour-effect 

is ignored, and only the ridge-like arrangement of the hair on the face and 
muzzle in parallel fringes is shown. 

which converge and stand up so as to form a well-marked raised 
stripe, “fringe” or “ridge” *, and intermediate tracts of smooth 
flat-lying hair. In some of the Museum specimens the more 
crowded upstanding hairs appear to be coloured more darkly than 
those of the intermediate tracts, but really the colour-effect is 
due to the pigment of the free ends of the hairs showing, whilst 
the thicker cortical substance of the bases of the hairs is white 

* The ridge of upstanding hair corresponds, it must be noted, to what isa shallow 
wrinkle-like groove in the feetal skin. 


wth . 

pie aN 
rR oe PP. 

th co pe 
an 4 ae 4 
iif INN 


Colour-striping (very slightly developed between eye and nostril) of the face of a 
Somali-land Giraffe (Giraffa reticulatus). Drawn from a specimen in the 
British Museum. 

(ile, Moron 

Z) al 


( yp ) AK x 
La, Le Sit Ha 

7 &. \ nc 
\ “i WY 

Hie iy 
/ Yi; ty 
Wy Yh 

TATE iif YY Yi if fh 
a Wi) Hii M/1, HT Wg Lyd) Hi 
Hi Hi Ha hil fii Ui Wy Hd yi'f WK, iy 

Walt) HI Wh WH. iol 
The same specimen as that shown in text-fig. 47, but with all colour omitted. The 

parallel bands between eye and nostril are not marked out in colour, but are 
merely elevated fringes or ridges of hair traversing a colourless region. 


(text-fig. 41, p.119). In one Somali specimen (@. reticulatus) the 
hairs are entirely white over a large part of the region of the face 
where the horizontal bands are developed. Yet these horizontal 
bands show up very distinctly on account of the more erect setting 
of the hairs (see text-figs. 47 & 48). In some cases the hairs of 
these ridges seem to have yielded more readily to destructive 
processes connected with taxidermy than have the hairs of the 
neighbouring tracts, and consequently the bands are marked out 
by nearly bald furrows or pathways. 

Another point of interest is that one of the horizontal pre- 
orbital hair-bands in the adult Giraffe directly leads up to the 
pre-orbital hair-whorl, and that the position of this hair-whorl 
appears to be farther in front of the eye in the Somali G. reticu- 
latus than in other Giraffes, whilst undoubtedly the hair is more 
erect and strongly developed on those ridges and on the hair- 
whorl in that species than in the other Giraffes which I have 
been able to examine. 

It would be interesting to ascertain how far the varying 
development of these colour-bands and parallel hair-ridges on the 
face of the Giraffe is constant in the different local varieties of 
Giraffe which have been distinguished. This isnot a matter with 
which I am at present able to deal. The purpose of the present 
communication is to call the attention of zoologists to a banded 
structure of the integument of the face in Giraffes which results 
in the appearance of dark and light parallel bands of hair and in 
the formation of strongly-marked parallel fringes or ridges of 
erect hair separated by bands of recumbent hair. These struc- 
tures are recorded in a foetal and in adult Giraffes, and appear to 
have hitherto escaped attention. 

The production of optically effective colour-bands by the mere 
crowding and direction of hairs along certain lines, without actual 
difference :n the pigmentation of the hairs of the lighter and 
darker stripes, has possibly some significance in regard to the 
origin and development of the more definite striping of the 
mammalian pelage so frequently shown, when dark and light 
stripes distinguished from one another are caused by the actual 
presence of pigment in the hair of the dark stripes, and its 
absence in the hair of the lighter or white stripes. 


Coloured drawing of the head and face, seen from above, of the foetus of a 
Giraffe removed from its mother which died in the Society’s menagerie in April 

The head of the foetus measured seven inches and one quarter from the anterior 
angle of the base of the outer ear to the extreme border of the upper lips. 

The specimen had been preserved in alcohol for six months. 

Details concerning the parentage &c. of this foetus are given by Mr. Beddard in 
Proc. Zool. Soc. 1906, p. 626. 

126 PROF, E, RAY LANKESTER ON [ Feb. 5, 

3. On the Hxistence of Rudimentary Antlers in the Okapi. 
By E. Ray Lanxester, M.A., D.Sc., LL.D., F.B.S., 
F.Z.8., Director of the British Museum (Natural His- 

| Received February 5, 1907. | 

(Plates VI. & VII.* and Text-figures 49-55.) 

We know a great deal more as to the horns of the Okapi than 
was the case when I communicated my description of that animal 
to the Society in 1901, and founded the genus Okapia. 

The two skulls sent home by Sir Harry Johnston—the first 
seen in Europe—were hornless, and it was at first a matter of 
doubt as to whether the Okapi wasa hornless Giraffid, or whether 
the male possessed horns whilst these two skulls were the one 
immature and the other that of a hornless female. 

During the printing of my memoir additional specimens were 
received in Brussels, and were transmitted to Dr. Forsyth 
Major in London for study and description. I saw in Dr. Forsyth 
Major’s possession a fine adult Okapi skull which had a pair of 
well-developed bony cones rising each by a broad base from the 
frontals, of which they appeared to form part. No suture was 
visible. An outline of this skull was published in my memoir 
by kind permission of Dr. Forsyth Major. I also was able to 
examine and to mention the existence of a curious structure 
discovered by that gentleman in regard to these ossicusps ; and I 
described it in the following terms :—‘“‘ The fine bony cones 
three inches long, which have made their appearance in the 

Text-fig. 49. 

Drawing of a fore and aft section through the tip of the ossicone of an adult Okapi 
in the collection of the Museum of the Congo Independent State. The section 
and drawing were made by Dr. Forsyth Major. 

The section shows the penetration of transverse fissures from the surface into the 
interior of the horn-tip. 

a, dense ivory-like bone; 6, posteriorly-placed transverse fissure; c, more anterior 
transverse fissure (marking off and presumably about to cut off and detach an 
anterior seement or plate of bone as a rudimentary “ antler ”) 

Brussels skull, show no suture at their base, nor any indication 
of origin as separate cap-like structures. For all that one can see 
they may be direct outgrowths of the frontal bone itself. Curiously 

* For explanation of the Plates, see p. 134. 

ici LON eae anvils 

Left. | Right. 
Left. Right. 


H.Grénvold dei London Stereoscopic Co imp- 

PLOT N= ene S Ol Oi i 

eas LOO, see valle 

G.M Woodward del. London Stereoscopic Co imp. 


ie Pal 
it Tne 

TRC ALTA ts: ue Als 
‘ 4, a pert v 

ow her) by 


enough, the point and posterior margin of the bony cone are 
polished as though it had protruded through the skin like a 
cervine antler. The point is separated by a suture from the rest 
of the ‘ ossicone,’ forming a small terminal cap of bone a third of 
an inch in depth. This curious structure, as well as a possible 
second suture a little lower down the ossicone, was pointed out to 
me by Dr. Forsyth Major. These appearances will be figured in 
that gentleman’s memoir on the Brussels’ specimens.” This is 
the first and so far the only published notice of the antler-like 
tips of the Okapi’s horns. The figure prepared by Dr. Major of 
the section made by him through the end of this Okapi’s ossicone 
is reproduced in the text-figure here appended (text-fig. 49). 
Dr. Major does not himself propose to publish anything further 
at present on the Okapi, and the little drawing has been placed 
in my hands by him. A tracing of it was also kindly sent to me 
by M. Fraipont, of Liége. 

The further history of our knowledge of the horns of the 
Okapi has been complicated by the arrival in Europe of various 
specimens, concerning the sex of which either erroneous informa- 
tion or none at all has been given by the natives from whom the 
specimens were obtained. Thus Dr. Forsyth Major was led to 
suppose that the female Okapi has a small unattached ossicone, 
some two inches in length, when adult, but he subsequently came 
to the conclusion that this supposed female was in reality a young 
male. In ‘ La Belgique Coloniale,’ No. 21, May 1902, Dr. Forsyth 
Major wrote :—“ L’Okapi posséde deux cornes frontales, recouvertes 
dune peau velue, plus petites, de forme conique et presque verti- 
cales chez la femelle; plus grandes, dirigées obliquement en 
arriére et en peu triangulaires chez le male.” 

At a subsequent date Dr. Forsyth Major came to the conclu- 
sion that the specimen supposed to be a female possessing small 
j * ossicones, was in reality a young male (Proc. Zool. Soe. 

». 339), and that the female Okapi is hornless, whilst the 
ie possesses “horns” which make their appearance as 
conical structures, ossifying independently of the sub- 
nes (as in the Giraffe) and becoming firmly ankylosed 
mtal bone in the adult—a boss-like upgrowth of which 
0 the structure of the complete horn. 

s little room for doubt that this is the true account of 
r, though we still are in want of full information as to 
icters of the adult female Okapi*. In a subsequent 
vation I shall be able to give more precisely the characters 
» types of skull, supposed to be that of the horn-bearing 
the hornless female, respectively. The skulls carrying 
sed or merely loosely-attached bony cone on the frontals 

‘0 the uncertainty which exists as to the origin of skin and skull which 

»s sent home from Africa as belonging to one individual, whereas they 

in certain cases belong to distinct individuals, it is still doubtful as to 
wuetner tne female Okapi has or has not in the adult condition a small knob-like 
protuberance of the integument, separable from the subjacent bone and representing 
the horn of the male. 

128 PROF, E. RAY LANKESTER ON [ Feb. 5, 

(supposed to be those of males) are longer and narrower than the 
equally large or larger skulls devoid of any bony cones in connec- 
tion with the frontals (supposed to be those of females) *. What- 
ever opinion is held, or whatever decision may be ultimately 
arrived at in regard to these two types of skull, it is the fact that 
they are very distinct from one another and that all the Okapi skulls 
which I have examined can be definitely assigned to one or the 

Text-fig. 50. 


Rudimentary free ossicone of hemispherical shape from the skin overlying the frontal 
bossed region of the skull of an Okapi of the broad-skulled type—sub-adult 
(deciduous molars very much worn, premolars not yet visible ; third lower molar 
in use on both sides, fifth cusp shows slight wear). 

a, natural size; 6, enlarged. 

Text-fig. 51. 

Section of the ossicusp drawn in text-fig. 50, to show the incomplete ossification. 

other of these two types. There is no third form known. The 
two types may perhaps be best distinguished as O. johrestoni (the 
name I gave to the broad hornless sub-adult skull accompanying 

* One of these broad-skulled specimens has, however, been found to possess a pair 
of completely detached bony ossicones of minute size embedded in the integument. 
The specimen is a little older (as indicated by the dentition) than Sir Harry John- 
ston’s larger individual (that mounted in the British Museum), but is not quite 
adult. It belongs to Messrs. Rowland Ward. I give here figures of the minute 
ossicone (text-figs. 50 & 51). 


the skin sent home by Sir Harry Johnston) and 0. liebrechtsi, 
the name given by Dr. Forsyth Major to the more elongate and 
narrow type of skull, which is that usually provided with bony 
cones attached to or ankylosed with the frontal bones. It is 
important to note that Dr. Major figures a skull (Proc. Zool. 
Soe. loc. cit. p. 423) which is hornless and is regarded by him 
as that of a female of the elongate type, O. liebrechtsi. I hope 
shortly to publish some measurements and outlines of these two 
types of skull. I have examined three of the UO. johnstoni-type, 
and five of the O. liebrechtsi-type. Though there is considerable 
variation in the number and breadth of the white stripes on the 
fore and hind limbs of the skins of Okapi received in this country 
(including the excellent specimens obtained independently by 
Major Powell Cotton and by Captain Boyd Alexander from widely 
separated localities, the former from the Ituri Forest, the latter 
from the Welle River), I have seen no evidence that a different 
striping of the skin is associated with the difference of skull-form. 
On the contrary, there is positive evidence that the striping of 
the skin is very nearly identical (though no two specimens are 
exactly alike) in animals which possessed the liebrechtsi form of skull 
with that exhibited by the mounted specimen (0. johnstoni) with 
hornless skull, sent home by Sir Harry Johnston, figured by me, 
and now in the British Museum. Nevertheless, it is true that 
direct and convincing evidence is as yet wanting for the conclusion 
that O. liebrechtsi is merely the male of O. johnstoni. 

When I had an opportunity (in 1904) of examining the fine 
skin of the adult (supposed) male Okapi, presented by the Congo 
State to the Museum of Paris, which is set up in the publie 
gallery there, I was especially anxious to note the state of the 
horn-tips. I found that they were represented in the mounted 
specimen and were seen projecting through the skin which 
clothed the ‘‘ossicone” up to a limit of about half-an-inch from 
the tip. From this level the dense bony matter was naked. 
It showed in each horn two fine transverse grooves, as 10 
the ossicone examined and sliced by Dr. Forsyth Major. This 
went far to prove that the condition noted by him was not 
exceptional or morbid, and accordingly I have examined the 
ossicones of other specimens of adult male Okapis, as opportunity 
occurred. Several skins and imperfect skeletons have been 
received in London by dealers in zoological specimens, and I am 
especially indebted to Messrs. Rowland Ward & Co. for the 
opportunity of examining the ossicones of four adult Okapis. 
Of two of these individuals I have had the ossicones drawn 
(PI. VI.) so as to show the free termination from different points 
of view. The two other specimens examined by me presented 
the same remarkable appearances as those figured, and as shown 
by the Paris Okapi, but I was unable to procure carefully drawn 
figures of them. Thus, including the Brussels skull examined 
by Dr. Forsyth Major, I have ascertained the existence of these 
transverse grooves or fissures in six adult male Okapis. I have 

Proc. Zoou. Soc.—1907, No. IX. 9 

130 PROF. E. RAY LANKESTER ON [ Feb. 5, 

also evidence of their existence in a plaster cast of another 
specimen which passed through the hands of Messrs. Rowland 
Ward & Co. 

An examination of the figures given in Plate VI. shows that 
in all four ossicones (the right and left of two adult male Okapis) 
the free terminal region is smooth and polished, forming a cap of 
about half an inch in length, whilst this region is followed by a 
rougher substance, furrowed on the surface. The polished region 
projects beyond the skin, the rougher region is clothed by the 
living integument. In all there are very deep horizontal fissures 
in the polished material of the “cap.” These fissures are some- 
what irregular in form, and it is impossible without making a 
section (which I had not permission to do)* through the solid 
material to ascertain their depth. They are of the same nature as 
those shown in the text-figure in section (text-figure 49, p. 126). 

I think there can be little doubt that these transverse fissures 
are caused by the ingrowth of the living tissue after the pro- 
trusion of the dense polished cap, so as to cut off the protruding 
portion and provide for its breaking off—Just as an antler is cut 
off and prepared for disruption in the Cervidee. A small conical 
piece is thus thrown off from the end of the horn or ossicusp, and 
may be regarded as a rudimentary or minute “antler.” 

But the process of discarding these minute points or antlers in 
the Okapi appears to differ from what occurs in the Cervide, not 
only in the minute size of the discarded segments, but in the fact 
that the preparation for the breaking off a second (and even a 
third) segment takes place before the first piece has been got rid 
of. The living tissue having absorbed the bony matter by a 
horizontal ingrowth and having created a transverse break in the 
continuity of the osseous substance (see text-figure), recedes for a 
distance of a sixth of an inch or less, and then again penetrates 
inwards, forming a new horizon of disruption; and from the 
appearance of the specimens figured in Plate VI., especially figs. 4 
and 5, it seems that this process of the recession of the living 
investment of the horn-tip and the subsequent ingrowth of the 
living tissue, may be again repeated before the most anterior 
piece is broken off, so that these horizontal fissures are visible on 
the surface of the horn-tip, following one another somewhat 

* Since reading this paper, I have been kindly permitted by the authorities of the 
Royal Scottish Museum to examine the horns of one of the specimens above referred 
to by making a section of the tip of the horn. The piece cut out has been drawn 
and then carefully replaced and cemented in position, so that no injury is done to the 
specimen. The skull lent to me by the Royal Scottish Museum is that of which I had 
already drawn the horn-tips in figs. 1 to 8, Plate VI., before it had passed from 
the possession of Messrs. Rowland Ward & Co. The sections drawn on an 
enlarged scale in text-figs. 52 & 53 explain themselves. It is seen that the grooves 
or fissures visible on the surface do not extend very deeply, but that there is evidence 
of resorptive activity in the form of certain branching canal-like structures lying 
deeply within the bony matter, which have probably been excavated by resorptive 
ingrowths from the soft surface tissues. 


Text-fig. 52. 


A. B. 

The diagram C shows the direction im which certain cuts have been made in the 

left osseous horn (ossicone or ossicusp) of the Edinburgh Okapi (also illustrated 
in P). VI. figs. 1 to 8). 
a, anterior border; b, posterior border.’ L, left side; R, right side. 

A isa drawing three times the natural size of the cut surface of the bisected horn, 

the bisection being effected in a plane erected on the line ab of the diagram C. 
It shows the eating in of the transverse fissures into the dense bony substance, 
and a number of irregular spaces and fissures which are probably cavities due to 
re-sorption of the bone. B is a drawing of the other half of the same bisected 

Text-fig. 53. 

Drawing of three times the natural size of the surface of a section obtained by 

cutting half of the same bisected horn-tip through a plane erected on the line R 
of the diagram, separating the shaded from the unshaded area. The penetration 
of the transverse fissures is shown. The cavities m, ”, 0 correspond to the 
transverse fissures labelled m, n, 0 in fig. 5 of Plate VI., representing the same 
specimen before it was divided. A 



We have no knowledge or indication that the shedding of these 
demarcated segments of the horn-tip is seasonal, nor indeed of 
the actual occurrence of such a shedding. However, the animal 
which is referred to in the explanation of Plate VI. as specimen A, 
shows in both right and left horn-tips a well-marked concavity, 
such as would correspond to the scar left by the “‘ shedding” or 
breaking off of a previously existing cap or segment of the dense 
bony substance. In fig. 2, this is seen in a view of the mediad 
(inner) face of the right ossicusp, and is marked w. In fig. 8 the 
same cavity is seen (and marked x) in a view taken from in 
front of, and somewhat above, the same horn-tip. 

It seems to me that we have in these appearances of the Okapi’s 
free or naked horn-tip (external termination of the ossicone) the 
evidence of a process of the same physiological significance as 
that seen in the seasonal removal of the antler in the Cervide. 
The continued contact of a deep tissue such as bone with the 
infective material of the outer world cannot be tolerated : necrotic 
organisms must effect a lodgment and gradually extend their 
ravages into the whole tract of the ossicone, and even to the 
bones of the skull. Accordingly, the exposed “tip” is cut off by 
a hone-absorbing ingrowth of the living tegumentary tissue, and 
the process of autotomy is active and recurrent. In the Okapi 
the process appears to be less elaborated and regularised than in 
the Cervide, and we do not know at present the details of its 
commencement or its final development. It is, however, certain 
that up to a late stage of growth, when the male Okapi is nearly 
of full size, the ossicone has not penetrated the integument with 
its tip, and that there is no indication of the polishing of the 
ossicone’s tip nor of transverse fissures caused by absorbent 
ingrowths of soft tissue. This is demonstrated by the ossicone of 
a male* Okapi of nearly full growth (the last molar of the upper 
series being not yet in use and the premolars only recently 
having superseded their deciduous fore-runners), which is illus- 
trated in Plate VII. This specimen belongs to a very perfect 
skeleton obtained, together with the skin, by Major Powell 
Cotton in the Ituri Forest, which is now in the British Museum 
(Natural History) at Cromwell Road. The ossicone figured is 
that of the right side. It is larger than that of the left side, 
weighing 31:45 grammes as against 28°15 grammes scaled by the 
left ossicusp. This asymmetry of the ossicusps of the Okapi, and 
a difference in the direction of the slope of these structures when 
right and left sides are compared, is to be observed in skulls of 
adult male specimens, and was mentioned by me in my memoir 
of 1901. 

The ossicones in Major Powell Cotton’s specimen have not yet 

* Major Powell Cotton ascertained that this specimen is a male, by an examination 
of the genital organs, and the skin prepared under his direction retains the external 


ankylosed with the frontal bone, but show an expanded base wath 
radiating trabecular structure of the bony material (see fig. 2, 
PV Uh: ys a peculiarity of surface which is repeated by the enlarged 
area of frontal bone upon which they rest—a condition which 
occurs also in young Giraffes. As shown by the drawings in 
Plate VIIL., there is no evidence in this ossicone of polishing or 
sharpening of the apex. The rough longitudinal grooves and 
furrows are continued to that region, and there are no transverse 
fissures. A vertical section (fig. 7, Plate VII.) of the apical region 
shows a very dense bony structure, but no trace of ingrowths 
from the surface. We may take it that this ossicone was still 
entirely covered in by the vascular living integument, as is the 
ossicone in the Giraffe throughout life. It furnishes us with 
a stage immediately antecedent to the fixation of the ossicone by 
ankylosis to the frontal bone, and antecedent to the breaking 
through of the integument by the apex of the bony cone. 

We may imagine the subsequent stages in the history which 
connect this specimen with those figured in Plate VI. The rubbing, 
polishing, and pointing of the ivory-like apex by use, the first 
horizontal ingrowth of the lacerated investing integument, the 
recession of that living investment after having established one 
horizontal discontinuity or plane of disruption of the dense bone, 
and the subsequent invasive ingrowth to form a second and by 
repeated recession and ingrowth a third such plane of disruption, 
and probably yet others. 

In conclusion, I would poimt out that it is quite conceivable 
that this cutting off of a series of antler-like tips from the 
ossicone of the Okapi, is a process independently set up in this 
Girafiid animal, having a similar physiological explanation to 
that which applies to the similar process familiar to us in the 

The different views which may be entertained, in the present 
state of our knowledge of the facts of embryology and early growth, 
as to the inter- relations of the ossicones of the Giraffidee, the 
antlers of the Cervide, and the bony horn-cores of the Bovide, 
are briefly stated in the preceding memoir on the “‘ Origin of the 
Lateral Horns of the Giraffe,” and to this I would now refer the 

[April 1907.—I add here text-figures (54 & 55, p. 134) of the 
ossicone of the specimen of Okapi brought home by Capt. Boyd 
Alexander from the Welle River and presented by him to the 
National Collection. There are many interesting features about 
the skin and skull which we owe to Capt. Boyd Alexander, and 
these I hope to describe hereafter. The horn or ossicone is that 
of an animal a very little younger than Major Powell Cotton’s 
(as inferred from the dentition). It is intermediate in size 
between that drawn in text-fig. 50 and that figured in Plate VII. 
Its bony substance is much less dense and ivory-like than that of 


Text-fig. 54. Text-fig. 55. 

Text-fig. 54.—Drawing of the right ossicone (not yet ankylosed) of the Okapi 

brought by Capt. Boyd Alexander from the Welle River. 

The relative age of this Okapi is indicated by the fact that whilst the deciduous 
molars are still in place in every socket, yet they are very much worn and all 
the true molars in both upper and lower jaw are “through ” and moderately 

Text-fig. 55.—Section through the tip of the same specimen to show the incomplete 
trabeculated ossification, contrasting with the dense ivory-like ossification of the 
ossicone of Major Cotton’s specimen (Plate VII.). 

The relative age of this Okapi is shown by the fact that the deciduous molars are all 
shed and the premolars moderately worn, whilst the molars are all “ through ” 
and moderately worn. 

the larger and older ossicone, whilst less spongy than that of the 
smaller and younger one. | 

Prater VI. 

Figs. 1 to 8 represent various views of the “horn-tips” of the ossicones or 
ossicusps of the Okapi in the Royal Scottish Museum, Edinburgh, of 
the natural size (Specimen A). 

Figs. 1, 4, 5, and 7 are views of the left horn-tip. 
Figs. 2, 3, 6, and 8 are views of the right horn-tip. 

Fig. 1. Left ossicone, outer side; y, depression due to shedding of a piece of the 

Right ossicone, inner (mediad) side; x, cup-like cavity due to shedding of a 
portion of the bone. 

. Right ossicone, outer side. 

. Left ossicone, inner (mediad) side; y as in fig. 1. 

. Left ossicone, from behind and below. 

. Right ossicone, from behind and below. 

Left ossicone, from in front and above. 

Right ossicone, from in front and above; «x asin fig. 2. 

Figs. 9 to 12 represent two views of the “horn-tips” of the ossicones of a 
specimen of Okapi lent by Messrs. Rowland Ward and now in 
America (Specimen B). 
Fig. 9. Left ossicone, outer side. 
10. Right ossicone, outer side. 
11. Left ossicone, from in front and above. 
12. Right ossicone, from in front and above. 


WADE oo 


Prate VII. 

The figures illustrate the ossicone of the right side of Major Powell Cotton’s 

male Okapi. 

. The specimen of the natural size, seen from the right. 

. The base of the ossicone, held by soft tissue to a similarly madrepore-like 
surface of the frontal bone. Natural size. 

3. Surface of the horn-tip, left side, enlarged to twice the natural size to show 
the absence of polishing and of transverse fissures. The ossicone was com- 
pletely covered by integument ; it had not been “ cut ” or emerged. 

. Similar view of the right side. 

. Similar view of the horn-tip from in front. 

. Similar view from behind. 

The horn-tip has been sawn through so as to remove the right-hand moiety 
of the tip. The extremely dense, ivory-like character of the bone of this 
region is thus demonstrated, and the absence of horizontal or other pene- 
as fissures (compare and contrast with the text-figures, especially text- 

g. 52). 



4. Description of Hyla resinifictriv Goeldi, a new Amazonian 
Tree-Frog peculiar for its Breeding-habits. By Prof. 
Dr. Emit A. Gorip1, C.M.Z.8., Director of the Para 

[Received January 21, 1907. | 

(Text-figures 56-59.) 

In its warty skin this remarkably fine Tree-Frog resembles Hyla 
tuberculosa Giinther, Hyla tawrina Steindachner, and Hyla venu- 
losa Laur. It is most closely related to the last. 

Length of a male, 8 cm. from snout to vent. 

Head semicircular. Space between the two nostrils slightly 
concave. Nasal region descending abruptly to the border of the 
mouth, almost at right angles to the frontal plane. Canthus 
rostralis running in a curve, rounded off. Nostrils, seen from 
above, forming slight prominences. Choane large. YVomerine 
teeth in two rows, forming an angle pointing forwards. Tongue 

Tympanum very distinct, moderate sized, rather smaller than 
the eye. Inthe male distinctly prominent vocal sacs between the 
rictus of the mouth and the insertion of fore-legs. 

Fingers III, IV, and V connected about half their length by 
a web; no perceptible rudiment of thumb. Finger-disks large, a 
little smaller than the tympanum. Outer border of arms without 
folds of the skin. When the hind legs are stretched forward, the 
tibio-tarsal articulation reaches the eye or the border of the mouth. 
Heels without appendage. 

Tubercles of sole of foot not so prominent as in Hyla taurina. 

Colour greenish-yellow with blackish-brown markings: a brown 
trapeze-shaped field between nose and anterior border of eyes; a 
light-coloured, broad band running from one eye to the other, 
the width of the eye, anterior bordering line straight, posterior 
line slightly curved backwards; a large dark field covering all the 
dorsal region, laterally rmmning down to the insertion of the fore 

136 DR. E. A. GOELDI ON A [ Feb. Ds 

legs, posteriorly leaving in the sacral region a triangular light- 
coloured space. Between the insertion of the hind legs, situated 
in the median line, there may be a small dark ring with a central 
dark point (text-fig. 57 A). Dorsal area either entire or laterally 
constricted about the middle (as in a fourth specimen not figured), 
or divided in two isolated parts by a light-coloured cross-band 
(text-figs. 56 and 57 B). The dark dorsal blotches are thickly 
studded with thorn-like elevations (pointed warts) with a light- 
coloured apex (text-fig. 56), more thickly agglomerated in the 

Text-fig. 56. 

Hyla resinifictrix, male (nat. size). 

parotoid region (left side 13, right side 22, mm one individual, 
and also in the posterior part 22+ 24 in one individual), besides 
an abundance of smaller elevations and diminutive granulations. 
A brown band around the upper arm, another around the fore- 
arm, very well pronounced (with some lightish spots, imitating 
the light central wart-spots of the back, but not raised); across 
the hand several narrower bands. Legs: thigh with two bands; 
lower leg with two broader bands, with the same whitish spots as 
above mentioned ; across the ankle and foot several narrower bands. 

Forehead with some dark, roundish spots. 

Upper border of mouth dark-marbled. 


Ventral surface light greenish-yellow, granulated and turning 
warty about the chest. Upper-arm band on the anterior side 
with two or three light spots; some light yellow elongated spots 
at the anterior border of the vocal sacs. Border of lower lip 
as well as the finger-disks of a delicate green. 

From a side view with the extremities folded, on the anterior 
extremity the dark bands on upper and lower arms appear to be 
continuous with the posterior border of the anterior large dark 
dorsal field; just as in the posterior extremity the dark bands 
nearest to the knee (femoro-tibial articulation) coicide with the 
posterior border of the second large dorsal field, while the bands 
nearest the heel (tibio-tarsal) coincide with the round sacral spot 
(text-fig. 57 A), the central spot forming with the lateral bands 
and dots a fan-like arrangement (confer also text-fig. 57 B). 

Text-fig. 57. 

AX< B. 

Hyla resinifictrix (diagrammatic). 

Comparison of the four specimens available shows a considerable 
extent of individual variation in the details of ornamentation. 
However, it is easily seen that certain general features persist 
fairly well, the most salient among them certainly being the 
broad light transverse band on the head, running from one eye 
to the other. 

The iris is golden, with a horizontal and a vertical black bar, 
forming a cross*. 

This large and strikingly coloured Tree-Frog presents a most 
curious novelty in its breeding-habits. Some years ago Mr, 
Boulenger, of the British Museum, published in an interesting 

* As described in H. venulosa by Boulenger, Ann. & Mag. N. H. ser. 5, vol. x. 
p. 327 (1882). 

138 DR. E. A. GOELDI ON A [ Feb. 5, 

article a tabular synopsis showing the different incubation-habits 
among the Batrachia. In this synopsis a unique position is held 
by the tree-frog Hyla palmata, which, according to observations 
made by me in the Serra dos Orgads (Rio de Janeiro), and 
published in the ‘Proceedings’ of the Zoological Society, 1895, 
forms breeding-bowls of the mud in shallow ponds. 

Text-fig. 58. 

Breeding-basin of Hyla resinifictrix, (A) side view and (B) section 

A still more peculiar eccentricity is presented by this beautiful 
new Amazonian tree-frog, Hyla resinifictrix. Inhabiting the 
virgin forest, it chooses certain tall trees for its dwelling, where 1t 
takes possession of a hollow branch (text-fig. 58, A and B), and 
constructs there as a nursery a good-sized basin of resinous sub- 
stances, with a central depression (text-fig. 59). As is well known, 


water and other liquids are preserved fresh in vessels varnished 
with pitch, and in like manner the rain-water which fills these 
resinous breeding-bowls presents excellent conditions for the 
hatching and development of the eggs and tadpoles, such as shade 
and freshness of water without contamination of decayed wood. 
Without having had as yet the good fortune to verify it by direct 
observation, I have abundant reason to suppose that the develop- 
ment stage of the tadpoles is exceedingly brief, analogous with 
the case of Hyla goeldii Boulenger in the Serra dos Orgads, Zyla 
venulosa in Para, and others. 

Text-fig. 59. 

Breeding-basin of Hyla resinifictrix, seen from above (from a photograph). 

One very interesting feature is the fact that this Amazonian 
tree-frog goes in search of the material with which to build the 
basin, and chooses for the purpose odorous resins which drop from 
the bark of certain trees, such as the aromatic “ breo-branco ” 
(Protium heptaphyllum) and others. 

Although the resin of the “cunnuart” is well-known to the 
Indians and mixed races in the Amazonian valley, constituting a 
commodity much sought for and of high price, the tree-frog 
itself was entirely unknown to all except the genuine forest- 
dwelling Indians. In spite of strenuous efforts it took me more 
than ten years to get on the track of this most mysterious 
Batrachian, and if finally my efforts were crowned with success, 
it was largely due to the friendly aid of the Tembé Indians at the 
Mission of Santo Antonio do Prata, at the River Maracana (interior 
of the State of Para), by the kind interest of Frei Daniel de 


Samarate, Director of the Mission. With their help I succeeded 
in obtaining several individuals, one of which ( 3) at least is still 
alive after spending nearly two years in a terrarium properly 
fitted up for it. Last year it gave me frequent opportunities to 
hear its voice, which is surprisingly strong, and sounds as “ queng- 
queng” three or four times repeated. 

The local name ‘ cunnuart,” evidently onomatopeeic, is formed 
by contraction of two Indian words ‘‘cunha=wife” and “art= 
toad”; the Indians say that this tree-frog always calls for the 
female when the moon shines. 

5. The Duke of Bedford’s Zoological Exploration in Hastern 
Asia.—III. On Mammals obtained by Mr. M. P. 
Anderson in the Philippine Islands. By OLDFIELD 
Tomas, F.R.S., F.Z.8.* 

[Received February 5, 1907. | 

In the early part of last year, after making the Korean 
Collection described in a previous volume of our Proceedings *, 
Mr. Malcolm Anderson paid a short visit to the Philippines, but 
was unfortunately attacked by fever, and after a gallant attempt 
to fulfil the object of his trip, was compelled to return to more 
northern and healthier latitudes. 

The chief object of Mr. Anderson’s visit to the Philippines was 
to obtain series of the interesting mammals discovered in 
Mindanao by Dr. E. A. Mearns, as the mountain fauna of this 
island was only represented in our National Nuseum by the 
duplicates from Dr. Mearns’s collection which the authorities of 
the United States National Museum had been good enough to 
send us. But these of course did not include any of the various 
new genera and species which had been described by Dr. Mearns 
on single specimens or on small series, and we therefore hoped 
that Mr. Anderson might be able to obtain some of them for us. 

Owing to Mr. Anderson’s illness the collection is quite small, 
only consisting of 16 specimens belonging to 6 species, but one of 
these proves to be new, while all are of interest as fillmg up gaps 
in our series. 


3. 756, 763. 92. 753, 761, 762. Mount Apo, Mindanao, 

Tam very doubtful if this form is at most more than a local 
subspecies of U. everetti$, described from Zamboanga. The type 
of the latter had been skinned after preservation in spirit, and such 
slight colour differences as there are may be due to this cause. 

* [The complete account of the new species described in this communication appears 
here; but since the name and preliminary diagnosis were published in the 
‘ Abstract,’ the species is distinguished by the name being underlined.—Ep1Tor. | 

+ P.Z.S. 1906, p. 858. 

{ See Mearns, P. U.S. Nat. Mus. xxviii. p. 425, 1905. 

§ Tupaia everetti Thos. Ann. Mag. N. H. ser. 6, ix. p. 250, 1892. 


3. 764. Mount Apo, Mindanao. 3000’. 

3. Mus sp. 

2. 748, 750. North Central Mindanao. 1100’, 3000. 

@. 754. Mount Apo, Mindanao. 3000’. 

A vat of the ratéus group. Dr. Mearns describes no less than 
three species of this group, but does not seem to be aware that 
the presence of spines in the fur in these rats 1s a most variable 
character, depending in all on age, and in some on season, the 
spines being shed and regrown periodically. 

4, Mus spacosus Mearns. 

Bullimus (g.n.) bagobus Mearns, P. U.S. N. M. xxviii. p. 450, 

@. 755. Mount Apo, Mindanao. 3000’. 

I fail to see any sufficient reason for the creation of a special 
genus to contain this species. The small supplementary cusps on 
the lower molars, on which Dr. Mearns mainly founds the genus, 
are not only present in many Malayan species usually referred to 
Mus, but they are even quite well-marked, though small, in his 
own specimens of Mus albigularis Mearns, also from Mindanao. 
None of the other characters mentioned by him appears to me of 
generic importance. 

Tt is unfortunate that Dr. Mearns had not had experience of 
the difficulties of Murine dental characters before venturing to 
describe genera of this group. Had he had such experience I am 
sure he would not have described Bullimus, nor would he have 
based another genus (Limnomys) on a single specimen with teeth 
“too worn to furnish characters distinguishing them from J/us,” 
unless the other characters were of a far more striking nature 
than appears from his account. 

Mr. Anderson’s specimen of JZus bagobus, being a female, 
enables me to state that the mamme number 1—3=8, a formula 
only occurring to my knowledge in two other Hastern species 
of Mus. 

5. Mus vuntcant Mearns. 
@. 752. North Central Mindanao. 4000’. 

A member of the common and widely spread concolor-ephippium 


Aosine, 12, At WOOT. 1. D (Wdelos 124) 
3g. 751. North Central Mindanao. 3000’. 29 February, 
1906. B.M. No. Type. 

A species with the general appearance of a blackish Akodon of 


the A. caliginosus group, or perhaps more of a member of the 
subgenus Melanomys. 

Size rather larger than in C’. fallaaw of Luzon. Fur close and 
fine, but rather stiff owing to the admixture of a number of fine 
flattened spines; hairs of back about 6-7 mm. in length. General 
colour above blackish brown, nearest to bistre of Ridgway 
but darker; the under surface nearly ss dark as the upper, 
without any line of demarcation, Individually the hairs of the 
back are dull slaty basally, with buffy tips ; the spines slaty with 
black tips. Ears very short, closely haired, brown. Whole of 
limbs, including hands and feet, blackish brown. Tail short, very 
finely scaled (scales about 18 to the centimetre), finely haired, the 
hairs about 14 scales long ; uniformly blackish throughout. 

Skull less flattened than that of C. fallax, but this may be due 
to youth. The brain-case long, comparatively parallel-sided, in 
continuation of the broad interorbital region. Supraorbital edges 
with scarcely a trace of beading. Outer plate of zygoma-root cut 
back just as in C. fallaw. Palatal foramina short, narrow in front, 
broadened behind, not parallel-sided as in fallax. 

Molars rather like those of a very simple type of J/ws, without 
any supernumerary lateral cusps. M, and M, each with a small 
median posterior cusp as in C. fallax. 

Dimensions of the type, which is barely adult :— 

Head and body 98 mm.; tail 68; hind foot (s.u.) 25; ear 13. 

Skull—greatest length 28 mm.; basilar length 22 ; zygomatic 
breadth 14; length of nasals 9°2; interorbital breadth 5-6; 
breadth of brain-case 12°3; palatilar length 11; diastema 7; 
palatal foramina 3°8 x 2:2; length of upper molar series 4:1. 

Hab. and Type as above. 

The characters of this remarkable species are a great puzzle, 
and only add to the difficulty of assigning a proper position to 
the type of the genus, Crunomys fallax. The single specimen of 
the latter is very old, and in the worn-down state of the teeth it 
was not clear whether they were or were not of hydromyine 
structure. The teeth of the present animal are certainly not 
typically hydromyine, but rather murine, while at the same time 
it is possible that in wearing down they might acquire the slight 
resemblance to hydromyine teeth shown by C. fallax. This being 
the case, I have decided to describe the species, though with doubt, 
as a Orunomys, but until younger specimens of C. fallax, or older 
ones of the present form, are available for examination, it would 
be advisable not to express any definite opinions as to the 
systematic position of either. As a species C. melanius is at once 
distinguishable by its blackish colour and heavier feet. 

7. SUS sp. 
4 skulls. Mount Apo, Mindanao, 

P.Z.S. 1907. Pl. VIII. 

J.Green photo imp. 


P.Z.S. 1907. PI. IX. 

nt pati 

J.Green photo imp 





P.Z.S. 1907. PI. X. 

ae % 

ea FW al 
Oe RR S&S eS = 
2 SinbegNe Se 



; Wows 

J.Green photo Imp. 



February 19, 1907. 
Sir Eymunp G. Lover, Bt., Vice-President, in the Chair. 

The Secretary read the following report on the additions that 
had been made to the Society’s Menagerie in January 1907 :— 

The registered additions to the Society’s Menagerie during the 
month of January were 108 in number. Of these 55 were 
acquired by presentation and 11 by purchase, 41 were received 
on deposit, and 1 was born in the Gardens. The total number 
of departures during the same period, by death and removals, 
was 163. 

Amongst the additions special attention may be directed to :— 

An Agile Gibbon (//ylobates agilis) from Borneo, purchased on 
Jan. 17th. 

A Wild Cat (Felis catus) from Inverness-shire, presented by 
Mr. G. W. Henderson, F.Z.5., on Jan. 26th. 

Two Superb Tanagers (Calliste fastwosa), a Crowned Tanager 
(Tachyphowus coronatus), and two Scarlet Tanagers (Rhamphocelus 
brasilius), from Brazil, purchased on Jan. 21st. 

Dr. C. I. Forsyth Major, F.Z.8., exhibited remains of a Bear 
from the superficial deposits of a cavern in the mountains of 
Corsica, where Bears, though now extinct, were formerly nume- 
rous, at least up to the sixteenth century. Despite the fact that 
no truly fossil Bears were as yet known from Corsica, Dr. Forsyth 
Major considered the Corsican Bear to have been autochthonous, 
whilst in his opinion the recent Mammals of Corsica (and Sardinia) 
had been, almost without exception, introduced by human agency. 
In any case they could not be adduced as proofs of a recent 
connection of those islands with either of the nei ghbouring 

The following papers were read ;— 

1. On English Domestic Cats. By R. I. Pococn, F.L.S., 
F.Z.8., Superintendent of the Zoological Society’s 

[Received February 5, 1907.) 

(Plates VIIIT_—-X.* and Text-figure 60.) 

1. The Classification of English Domestic Cats. 
Domestic Cats in Great Britain are classified by the National 
Cat Club under two headings :—(1) Short-haired ; (2) Long- 
haired, otherwise called “Persians” or ‘“ Angoras,” The 
“ Persians” are subdivided according to colour into “ Blacks,” 

* Foy explanation of the Plates, see pp. 167, 168. 

144 MR. R. I. POCOCK ON [ Feb. 19, 

“¢ Smokes,” “Sables,” “‘ Blues,” “‘ Oranges,” “‘ Creams,” “¢ Whites,” 
“Tabbies,” ‘“ Tortoise-shells,” &¢. The ‘Short-haired” are 
similarly distinguished, after the elimination of certain types like 
“Siamese,” “ Manx,” and “ Abyssinians,” to which special classes 
are assigned *. 

From the exhibitors’ and breeders’ points of view this arrange- 
ment has much to recommend it, and probably supplies the most 
feasible and satisfactory method of classifying the various breeds 
that are set side by side for comparison. The animals must be 
sorted out upon some basis, and it would perhaps be impossible to 
suggest a substitute that would meet the requirements of fanciers 
equally well. 

But, from the standpoint of affinity and descent, I think it is 
open to the criticism that a primary impor fance is given to 
characters like the length and thickness of the fur, the tint of the 
ground-colour, and the absence of the tail, which can be shown to 
have no great systematic value; whereas a quite subsidiary signi- 
ficance is attached to the nature of the pattern of the stripes, a 
character which should be of the greatest moment in differen- 
tiating the breeds, if confidence be paced i in the analogy supplied 
by existing species of the genus Felis. 

It may fairly be asked, however, why a greater taxonomic value 
is claimed for the “pattern” than for the three characteristics 
mentioned above upon which fanciers establish their breeds. The 
answers are briefly these :— 

1. The shortness of the hair in tropical Leopards and Tigers as 
compared with those that come from colder countries shows that 
no great systematic importance can be attached to length of 
coat. The difference between the extremes of long- and short- 
haired Domestic Cats is admittedly much greater than that between 
long- and short-haired Leopards or Tigers: but all gradations 
between ‘“ Persian” and ordinary Cats exist, so that no hard-and- 
fast line can be drawn between them; and probably no one 
doubts that the luxuriant growth characteristic of the former 
breed has been preserved and increased by artificial selection. 
In the matter of coat, the adaptability of Domestic Cats to 
changed conditions is proved by the evolution of a thick-haired 
breed in the Pittsburgh refrigerators 7 and in the arctic island of 
St. Paul t, and by the alleged shortness and stiffness of the fur in 
Domestic Cats from Mombasa in Hast Africa §. 

2. The conclusion that the shortness or complete absence of the 
tail in so-called “ Manx Cats” has been brought about by selective 
breeding from favoured sports must be regarded as beyond 
dispute. There is no such thing as a tailless species of Felis 
amongst fossil or recent forms. The tail is short in Lynxes, but 

* ©Cats and all about them,’ 1902; and ‘The Book of the Cat,’ 1903, -by 
Frances Simpson. 

t Quoted from Lydekker, ‘Cats, &c.’ p. 159 (1896) (Allen’s Nat. Library). 

* See H. C. Marsh, ‘Darwinism,’ p. 21 (1883). 

§ Darwin, ‘ Animals and Plants under Domestication,’ p. 58 (ed. 1905). 


it cannot be claimed that an intermixture with any known 
species of Lynx has contributed to the abbreviation of the tail in 
the “ Manx.” When and where this breed arose has been much 
debated. Some have suggested Spain as its original country, 
others China. But in all probability the “ sport” has appeared 
independently and been preserved by selective crossing on many 
occasions and in many places. 

The liability of the tail to modification in Domestic Cats is 
shown by the frequency with which it is kinked or twisted as 
well as shortened in Bur mese, Siamese, and Malaccan specimens ; 
and by the stunting it acquired in a few generations in the Cats, 
above alluded to, that were bred in the frigid climate of St. Paul’s 
Island and in the cold-storage warehouses of Pittsburgh. 

3. With respect to tint or the ground-colour as a whole, irre- 
spective of pattern, it must be remembered that albinotic and 
melanotic “sports” may arise in almost any group of Mammals. 
Amongst the Cats, albinos appear to be rare in a state of nature, 
probably in part on account of a correlated inherent delicacy 1 in 
organisation, accompanied possibly by defective sight or hearing ; 
probably in part on account of the conspicuousness of the 
coloration making capture of prey and escape from enemies, 
especially during cubhood, difficult. There are one or two 
records, however, of albino Tigers—the species, be it noted, in 
which the young after leaving the mother’s protection are more 
capable of taking care of themselves and less liable to attack than 
any other Cat, with the possible exception of Lions. Melanisms 
are far more frequently met with in the genus. Black Leopards 
are familiar to all; black Jaguars are not uncommon, and black 
Tigers, Caffre Cats, and Servals have been recorded, 

In other species the tint may be dimorphic, dark grey or dark 
brown and red or chestnut examples being about equally abundant, 
as in the 8S. American Felis jaguarondi, the West African Velis 
aurata, the Oriental F’. temmincki, and the Bornean /. badia. 

Thus in the genus Felis colour-variation in a state of nature 
may depart from the normal in the direction of “black,” or 
“ved,” or “ white,’ without respect to locality. Moreover, 
geographical variation also attesting inherent instability of tint 
is met with. The greyness of the Persian Leopard as compared 
with the rich tawny yellow of South Indian specimens is a case 
in point. 

The tints of Domestic Cats themselves establish the same con- 
clusion. Between “ blacks” and “ whites” all intermediates seem 
to exist. ‘ Blues” are self-coloured slate or lavender-grey cats 
and appear to be merely stages in the direction of “blacks.” 
“Smokes ” are darkish grey or blackish cats with the basal part 
of the hair white. ‘“‘ Silvers” are the “‘ palest conceivable edition ” 
of Smokes. ‘ Reds,” “ Yellows,” and ‘‘Creams” are variations 
from the normal in the direction of ‘‘ Whites.” 

4. On the other hand, notwithstanding individual and local 
variations, the pattern formed by spots or stripes in existing 

Proc. Zoou. Soc.—1907, No. X. 10 

146 MR. R. I. POCOCK ON [Feb. 19, 

species of Felis is on the whole constant. Stripes may break up 
into spots or spots may run together to form stripes, or both spots 
and stripes may be altogether fugitive. Yet even in extreme 
and rare cases of this nature, the general “character” of the 
pattern, when detectable, remains the same, and there is abund- 
ance of evidence that the animals breed true to the local type. 
Nor, so far as I am aware, is there any reason to suppose that 
dimorphism in pattern ever occurs or has ever occurred im any 
species of the genus Felis. 

It is needless to say more in support of the contention that if a 
decided difference in the “ patterns” of Domestic Cats exists, it 
must be regarded as furnishing a surer basis for their classification 
than the length of the hair, the tint of the coat, or the stunting 
of the tail. 

It may also be claimed with assurance that the pattern supplies 
a more important clue to the ancestry of Domestic Cats than the 
features just mentioned. 

Probably no one will dispute that all breeds of Domestic Cats 
have been derived from one or more than one ancestral type that 
was marked with bands or spots. This opinion is supported by 
two considerations. The first is this: spotting or banding of one 
type or another characterises the great majority of the existing 
species of Cats, using this term in its broad sense as co-extensive 
with the genus Felis*. A few self-coloured Cats, like Lions, 
Pumas, Caracals, and others, exist ; but their descent from striped 
or spotted forms is attested in most cases by the presence on the 
cubs + of markings which are subsequently lost or by indications 
of them, especially on the legs or lower parts of the body, in the 

The second pertinent point is the prevalence of stripes forming 
a pattern of one kind or another, both in “ Persians,” “ Short- 
haired,” and “ Manx” breeds, and the difficulty breeders experience 
in eradicating these markings in their efforts to preserve a 
particular self-coloured type. Frequently at all events the so- 
called “ blotched ” pattern can be detected in certain lights even 
in “ Whites ” and “ Blacks,” the two varieties which stand at the 
extremes of the colour-mutations of the diverse domestic breeds. 

Assuming, then, that domestic breeds are descended from one or 
more than one striped or spotted species, we may safely set aside 
the self-coloured forms as derivatives and consider only the 
striped or spotted types in looking for the origin and for a reliable 
basis for the classification of Domestic Cats. 

Domestic Cats in which the markings on the skin form definite 
patterns are called comprehensively “ Tabbies.” Of “Tabby” 
Cats, as fanciers well know, there are two kinds, the “striped ” and 
the “blotched.” These are not, however, regarded as different 

* TI purposely ignore here the subgeneric divisions of Felis adopted in Trouessart’s 
‘Catalogue of Mammalia,’ because I cannot admit that these connote, at least in all 
cases, natural assemblages of species. 

+ Young Caracals are not spotted or striped, but resemble their parents. 


breeds. Nevertheless the patterns appear to be fundamentally 
distinct from each other in the sense that the differences between 
them are not differences of degree, but of kind. Without 
assuming the existence in the past of a number of intermediate 
stages which do not appear to exist in the present, it 1s impossible 
to reduce them to a common plan and it is difficult to see how 
one can have arisen from the other unless per saltwm*. 

One or the other of these patterns, when the pattern is traceable 
at all, may be seen in Cats of the short-haired, long-haired, or 
Manx breeds, whether the ground-colour be grey or red, or black 
or white, or any other tint. Neither at Cats’ Homes nor at Cat 
Shows nor in the streets have I seen one cat out of the hundreds 
observed in which the pattern could not be assigned at once to 
one or the other of these types. Here, then, is evidence for the 
existence, side by side, of two fundamentally distinct kinds of 
“‘ Domestic Cats.” These I propose to regard as species, trusting 
to the analogy supplied by wild forms of the genus felis and 
knowing no reason for thinking, much less proof of the fact, that 
one has been derived from the other, either as a sudden sport or 
by gradual modifications under the influence of selective breeding 
or by inter-breeding with any wild species of the genus. 

The two types are described in some detail below (see pp. 151- 
153). They may be very briefly diagnosed as follows :— 

a. Sides of the body, from the shoulder to the root of the tail, 
marked with narrow wavy vertical stripes which show 
a tendency, especially on the thighs, to break up into 
spots ; no broad latero-dorsal stripe ...... Striped “ Tabby.” 

b. Sides of the body marked with three usually obliquely 
longitudinal stripes forming the so-called “ spiral,” 
“horseshoe,” or ‘ circular” pattern of fanciers; a 
broad latero-dorsal stripe on each side of the narrow 
median spinal stripe ............ sdsroialalasic'os Blotched “‘ Tabby.” 

It is difficult to ascertain from the writings of earlier natura- 
lists whether they were familiar with these two types of Cats or 
not. There is certain evidence that the blotched or true “Tabby” 
Cat was domesticated in Sweden and known to Linnezus as early 
as 1746. And that the Striped Cat was also domesticated in 
Kurope at a still earlier date is proved, I think, beyond doubt by 
the figures published in Gesner’s Hist. Anim. p. 345 (1551), and in 
Johnston’s ‘Quadrupeds,’ pl. Ixxil. p. 126 (1657). If these and 
later post-Linnean authors distinguished the two Cats, it must be 
inferred that they attached no significance to the difference of 
pattern, but regarded it as of the same value as the difference of 
colour and as the asymmetrical blotching of piebald specimens. 
Pennant, for example, speaking of the Tame Cat, says that it 

* The differences between these two types are described and figured by Mr. T. 
S. Rope (‘ Zoologist,’ 1881, pp. 353-357). Perhaps the omission of this paper from 
the ‘ Zoological Record’ may explain in part the want of consideration the facts 
have received from most recent zoological writers on this subject. . 


148 MR. R. I. POCOCK ON [Feb. 19, 

differs only ‘‘in color and some other trifling accidents” from the 
Wild Cat (British Zoology, vol.i. p. 94, 1812), and Kerr describes 
the stripes on the sides of the Wild Cat as being “perpendicular 
or spiral” (Anim. Kingd. p. 152, 1792), thinking apparently that 
the blotched ‘‘ Tabby” was a domesticated form of the European 
Wild Cat and that the shape and direction of the stripes were 
subject to variation and of no particular moment. 

Even the writings of later authors leave them open to suspicion 
on this point. Gray, for example, pointed out that a Domestic Cat 
brought by Darwin from 8. America was remarkable for its striking 
likeness to the Caffre Cat. But there is nothing surprising 
in this if, as I suppose from the description, the Cat in question 
was merely a slightly aberrant example of the Striped “ Tabby” 
breed. This Gray could hardly have failed to detect had he 
been familiar with the Domestic Cats of London. Again, since 
Blanford and Mivart both regarded Indian specimens of the 
“Striped breed,” known as /’. torquata, as examples of a genuine 
wild species, it may be inferred that they were both ignorant of 
the fact that Cats inseparable from that type, as figured by Cuvier, 
might be seen any day in London, where and at the time when 
their volumes, below (p. 159, footnote) cited, were being written. 

Setting aside breeders and owners of “ Fancy Cats,” who could 
hardly be expected to appreciate the significance of the differences 
of pattern above described, although well aware of their existence, 
two scientific writers on Cats must be mentioned as clearly 
apprehending the fact. One of these is Rope, whose paper, 
published in 1881, has been already mentioned; the other is 
that astute observer Blyth, who so long ago as 1845* pointed 
out that the two types of pattern are found in the Domestic Cats 
of Europe. In an additional note on the subject published 
in 18567, he spoke of the true Tabby pattern as being possibly 
a “ modification (and a very remarkable one) of the markings of 
the wild F. sylvestris of Europe.” 

Rope apparently did not know of Blyth’s papers. All the 
more remarkable is it therefore that both authors describe this 
Tabby as being marked with pale streaks on a dark ground; and 
this description was repeated by Hamilton in his notice of Rope’s 
paper. It is obvious, however, that no Cat can be scientifically 
described as marked in this way. Hence it is possible that the mis- 
leading terminology employed by Blyth, Rope, and Hamilton may 
have suggested to the readers of their writings that the blotched 
“Tabby” markings belong to the same category as the piebald 
or skewbald, usually asymmetrical coloration of black-and-white 
or brown-and-white cats, dogs, horses, cattle, and other tamed or 
domestic mammals infected with either melanism, albinism, or 
erythrism, or with any two of these taints, or with the three 
combined. This may partly account for the fact that the difficulty 
of deriving the blotched Tabby pattern from that of /. ocreata or 

* Journ. As. Soc. Bengal, xiv. pt. 1. pp. 342-343. 
“+ Op. cit. xxv. p. 443. 


F. sylvestris, and of accounting for the origin of the breed, seems 
never to have been fully realised, or, if realised, never seriously 
faced. And doubtless a contributory cause to this result has 
been the artificial character of the classification of Domestic Cats 
adopted by fanciers, which gives emphasis to valueless features 
and obscures the fundamental importance of the pattern. 

2. On the Characters and Names of the Two Types of Domestic Cat 
and the Name of the European Wild Cat. 

In the foregoing and following pages the European Wild Cat 
is referred to as Felis sylvestris. This requires explanation, since 
the species in question has, by almost common consent, been 
hitherto called Felis catus. 

In my opinion there is no possibility of evading the conclusion 
that the Cat to which Linnzus gave the name catus was not the 
European Wild Cat, but the Domestic Cat of the blotched Tabby 
kind. In the 10th edition of the ‘Systema,’ 1758, accepted as the 
starting-point in zoological nomenclature, Felis catwus is character- 
ised in such terms as to leave no room for doubt on this head. In 
the first place, there is a back reference to the ‘ Fauna Suecica,’ 
1746, where of Felis catus it is said “habitat in domibus; cicurata: 
exotice originis.” In the second place, the diagnosis in the 
‘Systema’ runs as follows :—‘“ Felis caudata, elongata ; corpore 
fasctis nigricantibus ; dorsalibus longitudinalibus tribus ; laterali- 
bus spiralibus.” 'The spiral lateral stripes are obviously those that 
form the so-called “ circle” or “horseshoe” characteristic of the 
blotched ‘‘ Tabby ” ; and the three dorsal longitudinal stripes are 
also typical of that animal. This description is totally inapplicable 
to the European Wild Cat, which moreover does not occur in 
Scandinavia, and was apparently unknown to Linneus, except 
from books. It is equally inapplicable to the “Striped” race of 
Domestic Cats. On the other hand, it exactly fits the blotched 
Tabby Domestic Cat, and was quite likely taken from a specimen 
lying on Linneus’s hearthrug at the time. 

There is no doubt from the context and the bibliography in 
the 10th ed. of the ‘Systema’ that Linnzus believed this Cat 
and the European Wild Cat to be the same species of animal ; 
but that error does not in any way affect the certainty of the 
conclusion that he knew enly the domestic blotched “Tabby” and 
described it as catus. ; 

The name “ domestica” must, I think, be discarded as a 
synonym of caétus, since both Schreber and Erxleben, the two 
earliest post-Linnean authors to use that term, quote under the 
heading domestica Linneus’s diagnosis of catws. And since all 
the so-called Tortoise-shell Cats that I have seen belong to this 
type, the name hispanica Erxl. may also be placed amongst the 
synonyms of catus ; vulgaris Fischer is also a synonym. 

The name ‘catus,” then, is no longer admissible for the 
European Wild Cat. For this species there are two names 

150 MR. R. I. POCOCK ON [ Feb. 19, 

available, which were apparently published in the same year ; 
and between these a choice has to be made. These are :— 

ferus Erxleben, Syst. Regni Anim. i. pp. 518 & 522 (1777). 
sylvestris Schreber, Siug. ili. p. 397 (1777), pl. evii. a. (date 2)*. 

The synonymy of felis catus ferus published by Erxleben 
shows that he followed Linnzus in regarding the European Wild 
Cat as what may be called the “agriotype” of the various 
domestic breeds, such as domestica, angorensis, hispanica, and 
cerulea, described on pp. 520-522; and the excellent diagnosis 
of the Wild Cat published on p. 522 beginning Catus Yerus major, 
proves that he was acquainted with that species, and applied the 
name ferus to it, and not to feral examples of a domestic breed. 

But for the following reasons I think sylvestris should be 
preferred :—(1) Schreber adopted this name from Brisson’s Regn. 
Anim., Quadr. p. 265, which, being published in 1756, is pre- 
Linnean so far as nomenclature is concerned. Nevertheless, the 
diagnosis of /, sylvestris suggests that Brisson distinguished this 
Cat from the domestic catus of Linneus. (2) Erxleben quotes 
both Brisson’s and Schreber’s works in his _ bibliographical 
synonymy of /. catus ferus. Of Schreber’s work he correctly 
cites pl. evii. A. inscribed Felis Catus Linn. ferus. Hence I infer 
that he saw or knew by hearsay of this plate. I cannot, however, 
find proof that it was published, properly speaking, before the 
issue of the text in 1777. Since, therefore, the name sylvestris 
in the text has page priority over the name /erws on the plate, 
preference should be given to the former and the latter regarded 
asasynonym. (3) The name sylvestris has been used for this 
Cat by some modern writers, notably by Blyth and Hamilton. 

For these reasons I think it advisable to adopt sylvestris instead 
of ferus as the specific name for the European Wild Cat. 

Apart from the convenience of having a name for a Cat which 
not uncommonly occurs feral in the tropics, the determination of 
the correct name for the “Striped” Domestic Cat is a matter 
of no great moment. Moreover, it is practically impossible to 
settle with certainty which name should be chosen out of the 
many given by earlier writers to various domestic breeds in 
different parts of the worldt. In addition to domestica and 
hispanica, already referred to, the following are the most im- 
portant :—ceruwea Erxl. for the ‘“ Blue Cat,” which seems to be 
a pale or incompletely melanistic sport perhaps of the striped, 
perhaps of the blotched type; angorensis Gmel., for the long- 
haired breeds, which were no doubt originally of the “ Striped ” 
type; ruber id., for a red variety with a dark dorsal stripe ; 
sinensis Kerr, for a Chinese race alleged to have pendulous ears ; 

* Kor discussion of the dates relating to Schreber’s work see Mr. C. D. Sherborn’s 
paper, P. Z.S. 1891, pp. 588-589. 

+ These names have to be catalogued to prevent their inadmissible use in a 
different sense by later writers (see Sherborn, Index Anim. p. 187, 1902). 


aureus id, (=longiceps Bechst.), for a yellowish, short-legged, long- 
headed, sharp-nosed breed, said to inhabit New Spain ; mada- 
gascarensis id., for a Madagascar Cat with a twisted tail; striata 
Bechst., for a black-striped “ Cyprian” Cat, which is diagnosed 
as follows :—‘ Mit schwarzen Streifen auf hellem Grunde, die 
auf dem Riicken gerade, auf den Schenkeln aber gekrimmt 
sind” (Bechstein, Pennant’s ‘ Uebersicht, etc.’ ii. p. 679, 1800). 
In a general way this description applies to both types of Cats 
under discussion, and perhaps on the whole more clusely to the 
“blotched ” than to the “striped” breed. But if the “Cyprus ” 
Cat came originally from Cyprus, a conclusion by no means 
justified by the epithet, it belonged in all probability, as did 
the Angora Cat, to the “Striped” and not to the “ Blotched” 

But whatever opinion may be held with regard to angorensis 
and striata, there is no doubt that examples of this Cat, whether 
feral or not, furnished types for the following forms :— 

F. torquata F. Cuvier, Hist. Nat. Mamm. pl. 126 (1826), 
recorded from Nepaul, Bengal, &c. (see infra, p. 165). 

F. inconspicua Gray, Charlesworth’s Mag. N. H. p. 577 

Vip ee J. A.S. B. xv. p. 169 (1846), xvii. p. 247, 
xxii. p. 581, from Candahar. 

F. ocreata agria Bate, P. Z.8. 1906, p. 317, from Khania in 

The figure of F. torquata and the description of 2. huttoni 
leave no room for doubting the identity of the Cats; and the 
types of inconspicua and ocreata agria are in the British Museum 
and have been compared by me with English examples of Striped 

This type (torquata) may be described as follows :—Ground- 
‘colour typically iron-grey or yellow-grey. The four cephalic and 
cervical stripes sometimes distinct, sometimes indistinct. The ad- 
median cervical stripes not abbreviated on the anterior area of the 
nape; the external cervical stripes not diverging from the middle 
line on its posterior area. Dorsal area of body from the shoulders 
to the root of the tail darker than the sides, the pigment often 
resolvable into three narrow, almost contiguous stripes, two 
laterals more or less interrupted, and a more complete median 
which is usually continued down the middle line of the tail. On 
the summit of the shoulder the lateral stripes are frequently 
thicker and more heavily pigmented than they are posteriorly. 
From the dark dorsal area on to the belly pass a number of 
vertical wavy transverse stripes, which are usually more distinct 
on the thoracic than on the abdominal region, where, as also on 
the thighs, they are more or less broken up into spots, which may 

* See Temminck, Mon. Mamm. i. p. 128 (1827); and Fischer, Syn. Mamm. p. 208 

152 MR. R. I. POCOCK ON [ Feb. 19, 

or may not show signs of transverse lear arrangement. The neck 
and the shoulders may be striped or merely speckled. The fore 
and hind limbs are transversely barred; the former are typically 
black behind up to the wrist and the latter up to the hock. The 
tail also is transversely barred above, sometimes throughout its 
length, sometimes only in its distal portion, the tip being black. 

In specimens that I regard as typical of this Cat the pattern is 
very similar to that of the European Wild Cat (7. sylvestris). 
In others the transverse stripes break up into more or fewer, 
larger or smaller spots. In others the spots are evanescent, the 
fur being merely “ticked,” a gradation being traceable from the 
first or sylvestris-type to the last, which are, I believe, the so- 
called “ Abyssinian” and “Ticked” breeds of the fanciers. I 
have obtained all these types from the Cats’ Home in Camden 
Town, London, in the space of a few weeks, as well as a fine red 
variety of the spotted kind. With the exception of the sylvestris-. 
type, all these phases can be matched approximately in the series 
of skins of /’. ocreata preserved in the British Museum. 

Specimens which are intermediate between the ‘ spotted” and 
the “ ticked” types—those, that is to say, in which the spots are 
very small and closely set, giving a blackish appearance to the 
skin except on the legs and ventral surface where the stripes are 
apparent—deviate from the normal in exactly the same way as 
the two South-African Leopards described by Dr. Giinther* differ 
from normally spotted Leopards. 

In the British Museum Collection there are skins of this 
Domestic Cat from such widely separated localities as Crete, 
India, Madagascar, Celebes, and Mexico. The specimen from 
Celebes was shot on Bonthain Peak. It is noticeable that its. 
stripes and spots are browner than in typical Kuropean examples ; 
and in the skull the posterior portion of the nasals is more com- 
pressed und the interparietal crest is higher and traceable as a 
ridge all along the parietals. The narrowness of the nasals 
recalls that of the skulls of two Siamese Cats that I have seen 
(cf. infra, p. 163), and the height of the interparietal crest is 
paralleled in the skull of a rufescent Indian Cat + which formerly 
belonged to Mr. H. C. Brooke and was, I understand, a well- 
known prize-winner at Cat Shows some few years ago. For 
further information about torquata, see below, pp. 164-165. 

The following description, taken from a number of skins, will 
perhaps convey a tolerably clear conception of the plan (¢f. text- 
fig. 60, p. 154) to which the patterns of F. catus, however 
diversified in detail, are reducible :— 

Ground-colour typically brownish or grey, frequently washed or 
clouded with blackish. Head and face normally striped. ‘The 
internal or admedian cervical] stripes abbreviated on the fore part. 

* P.Z.S. 1885, p. 243, pl. xvi.; id. op. cit. 1886, p. 203, fig. See also W. L. 
Sclater, Mamm. S. Afr. i. p. 36, fig. 10 (1900). 
+ This Cat, Mr. Brooke tells me, was brought as a kitten from a hotel in Bombay. 


of the nape, and entering the anterior apex of a diamond-shaped 
space (c.sp.), which is bordered laterally by the posterior ends of the 
external cervical stripes and the anterior ends of a pair of broad 
suprascapular stripes ( The latter lie near the middle line 
on the summit of the shoulder, and diverge forwards to meet at 
an obtuse angle the posterior ends of the backwardly-diverging 
external cervical stripes. From this angle stripes pass obliquely 
forwards and downwards on to the sides of the neck and throat, 
the posterior of them lying in front of the shoulder and forming 
a throat collar. From the suprascapular stripe one broad vertical 
stripe, the scapular stripe (sc.), passes over the shoulder and loses 
itself in the uppermost of the transverse brachial stripes of the 
fore leg. This scapular stripe forms the anterior border of the 
postscapular space (, which is bounded behind by a second 
broad vertical stripe, the postscapular stripe (, descending 
from the posterior end of the suprascapular stripe. Extending 
along the middle of the spine from the posterior apex of the 
cervical space there is a very narrow spinal or median dorsal stripe 
(sp.), which passes down the middle line of the tail. Running 
forwards on each side of this from the root of the tail to the 
posterior end of the suprascapular stripe, which it frequently joins, 
there is a very broad latero-dorsal stripe (/at.dors.). Beneath or 
externally to this on the body there are three broad stripes 
whose direction is obliquely longitudinal. The upper of these, the 
supero-lateral (, forms a bold downward curve posteriorly, 
while the inferior, the infero-lateral (inflat.), curves upwards 
anteriorly. They thus partially circumscribe an elliptical or 
subcircular area, in which lies the short and broad medio-lateral 
stripe (, which frequently has the form of a large spot 
or blotch. These three stripes form the so-called “ horseshoe,” 
“ circular,” or “spiral” mark characteristic of this type of 
Domestic Cat. On the thighs there are broad transverse stripes, 
the upper of which, the femoral stripe, extends obliquely down- 
wards and forwards from a point beneath the posterior end of the 
latero-dorsal stripe to a point beneath the downcurved posterior 
end of the supero-lateral stripe. In the space between the three 
stripes just mentioned lies a large spot or stripe, which frequently 
fuses with the femoral stripe. On both fore and hind legs the 
stripes thin out and die away towards the paws, which are typi- 
cally black behind up to the wrist and hock. The tail is black at 
the tip, and marked throughout with transverse black bars, which 
are very broad where they touch the median caudal stripe. 

The pattern of this Cat varies considerably in detail with 
respect to the width and degree of fusion of the stripes. The 
cervical and postscapular spaces are sometimes hardly apparent, 
and the stripes may widen and fuse to such an extent that an 
almost totally black Cat results. It is possible that the blackness 
of some Cats is attributable to this process; but usually the 
blackness is due to the melanism of the ground-colour, the 
stripes retaining their normal width and being detectable under 

154 MR. R. I. POCOCK ON [Feb. 19, 

reflected light by the greater glossiness of the hair, The stripes 
in white Cats are often visible in the same way. Tortoise-shell 
variations of this Cat are nothing but partially erythristic or 
erythro-melanistic sports, frequently with pronounced albinism 

Text-fig. 60. 

_MED. LAT, ------------ 

SUP. LAT. -------------= 


Diagram of the Pattern of the Blotched Tabby (Felis catus). 

¢.sp., cervical space;, postscapular space; se., scapular stripe;, supra- 
scapular stripe;, postscapular stripe; sp., spinal stripe; lat.dors., latero- 
dorsal stripe;, supero-lateral stripe;, medio-lateral stripe ;, infero-lateral stripe. 

affecting particularly the legs, inferior areas, and even other parts 
of the body. In simple cases the blotching may be seen to be 
due to the red tint occurring only on portions of the stripes, 
the remaining portions of which retain their normal black hue. 


The result is a mixture composed of red and black stripes on a 
yellowish ground-colour. But in more complicated cases the red 
and black also invade the ground-colour in patches, either the 
red or the black, but more commonly the black, predominating. 
On these skins the original pattern is almost entirely obscured. 
It is a well-known but none the less singular fact that Tortoise- 
shell Cats are generally females and “ red” Cats generally males. 
This circumstance is the foundation for the saying that the ‘red ” 
Cat is the male of the “ Tortoise-shell ” ; but since the Tortoise- 
shell is as much melanistic as erythristic, male black cats have 
equal claims to be so regarded. 

3. On the Characters of the European and African Wild Cats 
(F. sylvestris and F. ocreata) and the Origin of the Domestic 
Types (torquata and catus). 

Criticism of the opinions of authors on the subject of the origin 
of “‘ Domestic Cats” must be prefaced by the remark that, with 
one or two exceptions, they failed to realise that an explanation 
was required of the origin of two distinct kinds of Cats differing 
so much from each other that no one would hesitate to regard 
them as representing widely divergent species if they occurred in 
a state of nature. 

A great deal has been written on this topic. Opinions at one 
time were nearly equally divided on the point, some authors 
regarding the European Wild Cat (/. sylvestris), others the 
Egyptian Cat (/’. ocreata) as the ancestral stock. Of late years, 
however, the latter species has grown greatly in favour for the 
distinction, partly on account of the unquestioned adoption and 
reiterated publication on the part of recent writers of a statement 
made many years ago, but none the less erroneous, that the 
Domestic Cat has a longer and more tapering tail than the Wild 
Cat of Europe*, and partly on account of Nehring’s assertion 7 that 
the Domestic and Egyptian Cats resemble each other, and differ 

* Macgillivray, in his excellent account of the British Wild Cat (‘British 
Quadrupeds,’ Jardine’s Naturalist’s Library, pp. 188-195, 1854), long ago pointed 
out that the tail of the Domestic Cat is thinner and more tapering than that of the 
Wild Cat, because its fur is much shorter. Dr. Hamilton also exposes the fallacy of 
the above-stated belief (‘ Wild Cat of Europe,’ pp. 41-42, 1896). One modern writer 
speaks of the tail of the Wild Cat as “clubbed.” I have never seen a specimen with 
a tail to which this epithet meaning broader at the tip than at the base could be 

+ SB. Ges. nat. Freunde Berlin, 1887, pp. 26-27, & ‘ Humboldt,’ 1888, pp. 189-141. 
Nehring (Zeits. fiir Ethnologie, xxi. pp. 558-9, 1889) suggests that Domestic Cats 
are descended from two main stocks—one from S.E. Asia, the other from N.E. 
Africa. From the former arose the ‘Chinese? Domestic Cat; from the latter the 
Egyptian House-Cat. The Egyptian House-Cat was the forerunner of the European 
Domestic Cat, with an infusion, especially in Germany, of the European Wild Cat. 
But whether the stock of the Asiatic House-Cat was F’. inconspicua Gray, or 
F. manul, or some other smaller Asiatic species, the author leaves undecided. It does 
not appear that Nehring realised the existence of the two types of European Domestic 
Cats I have described. Also he can scarcely have been acquainted with the 
peculiarities of F'. manul or with the nature of the affinity between F. incon- 
spicua and F. ucreata. 

156 MR. R. I. POCOCK ON [Feb. 19, 

from the Wild Cat in having the back of the hind Jeg from the hock 
to the pad blackish. This statement is negatived by three facts :— 
the area in question is not blackish in all Domestic Cats; nor in 
all examples of the Egyptian Cat; nor does it lack the dark tint in 
any specimens of the Kuropean Wild Cat that I have examined. 
The prevalent idea on this subject has been expressed as follows :— 
“The black sole of the foot suggests that the Caffre Cat is the 
chief stock from which the Domestic Cats of Kurope have been 
derived”; and “that the Kuropean Wild Cat was not the direct 
descendant |? lapsus for ancestor] of the domesticated breeds of 
the western part of the continent is rendered pretty evident by its 
short and clubbed tail, to say nothing of the absence of dark soles 
to the hind feet” *. And again:—‘It has been maintained 
by many naturalists that the European Domestic Cat is chiefly 
derived from the north-eastern race of this species [/’. ocreata] 
found in Egypt; at least, the domestic form is certainly not 
derived from the European Wild Cat” +. The foundation for this 
positive opinion entertained by Mr. Sclater may be found, I 
think, in his belief (Cat. Mamm. Indian Mus. ii. p. 934, 1891), 
taken from Blasius, that in the skulls of the Domestic Cat the 
frontal and squamosal bones are separated from one another by 
the parietals and alisphenoids, and the nasals are not produced 
posteriorly beyond the frontal processes of the maxille, the con- 
verse being the case in the Wild Cat. I have before me the | 
skulls of four ‘ London” Cats. In one of them the nasals project 
well beyond the maxille, in two as far back as the maxille, and in 
the fourth not so far. Again, the distance between the frontal and 
the squamosal is in one skull as much as 3 mm., in two others it is 
only about 1 mm. or less, while in the last the squamosal and 
frontal touch on one side but not quite on the other. Nevertheless, 
it is perfectly true that, as a very general rule, the squamosal 
and frontal are separated by the junction of the parietal and ali- 
sphenoid in Domestic Cats as they also arein J’. ocreata. But 
this is also the case in both the skulls of the Scotch Wild Cat that 
I possess=. Im one of them the parieto-alisphenoid bridge 
measures 4 mm., in the other 2mm. The bridge is also present 
in one of two skulls of this species in the Museum of the College 
of Surgeons, whereas in the other the frontal and squamosal have 
a long sutural union. 

Thus, as Dr. Hamilton has already shown, the claims of both 
Blasius and Nehring regarding the differences above alluded to 
between the Wild and Domestic Cats of Europe will not stand 
the test of the examination of skulls and skins. 

It appears to me that much barren discussion on this subject 
would have been saved by the realisation of the closeness of the 
affinity between the Egyptian and the European Wild Cats 

* “Cats &.’ in Allen’s Nat. Library, pp. 156 & 157 (tee), by R. Lydekker. 

+ ‘Mammals of S. Africa,’ i. pp. 43-44 (1900), by W. L. Sclater. 

{~ There is no reason to doubt that both these skulls belonged to pure-bred Wil 
Cats; this I can vouch for, since I have the skins. 


(Ff. ocreata and F. sylvestris). The type of pattern they present 
is not found in any other species of Felis, the nearest approach 
thereto being that of the Tiger (/. tigris) and in a remoter degree 
of Pallas’s Cat (7. manwl). The similarity in pattern between the 
two, coupled with their geographical distribution, almost induces 
the adoption of the view that they are but northern and southern 
forms of the same species. There is as yet, however, no positive 
evidence that they actually intergrade to the extent of justifying 
the conclusion that their distinctive features are merely of sub- 
specific value. 

The characters they have in common are as follows :— 

Close resemblance in the shape, size, and structure of the skull 
and teeth *. 

General similarity in size and shape of the body, of the paws 
and ears,and in the length of the tail, though this organ is 
apparently a little longer at least in North-African examples of 
F. ocreata than in F. sylvestris. 

General similarity in pattern: in both species the sides of the 
body are typically marked with wavy vertical dark stripes ex- 
tending from the spine to the belly, and better defined, as a rule, 
on the thoracic than on the abdominal portion ; the upper parts 
of both fore and hind limbs are transversely banded with broad 
bars, almost always darker than the transverse stripes on the 
body ; the fore leg is black below from the toes to the wrist and 
behind at the elbow, the two internal brachial stripes, normal in 
felines, being well developed ; the hind leg is normally blackish 
below from the toes up to the hock. The distal portion of the 
tail has a black tip and about three well-defined black stripes pre- 
ceding it, the proximal portion of the tail being much less distinctly 
barred and marked at best with an ill-defined median dorsal 
stripe. The throat is unstriped or only indistinctly striped, and 
usually exhibits a white spot ; the thoracic and anterior abdominal 
areas of the ventral surface are spotted ; the posterior abdominal 
and inguinal areas are unspotted and tinged with yellowish buff, 
especially on the inner side of the thighs. Identity prevails even 
in the colour of the individual hairs, which are slate-grey at the 
base, then buff or cinnamon, then black, the distal portion being 
yellowish or greyish white with a black tip. 

The principal differences between them are as follows :— 

in F. sylvestris the four paired stripes on the head and neck are 
well defined; on the occipital region they diverge from the middle 
line and run backwards almost to the shoulder as four wavy widely 

* Amongst the skulls in my own collection and in that of the British Museum I 
have so far failed to find any constant characters for distinguishing the two species. 
By the sum of a number of small features in the teeth and bones, the skull of 
sylvestris can generally be recognised from that of ocreata. But up to the present 
every character which I thought might prove to be distinctive of sylvestris has 
broken down under the examination of a long series of skulls of ocreata, a species 

‘which, in its broad sense, is extremely variable in its cranial osteology. So far as the 
skull is concerned, the differences between the two species must be regarded as of 
“ subspecific ” value. 

158 MR. R. I. POCOCK ON [ Feb. 19, 

separated stripes, the admedian pair being, as a rule at all events, 
better emphasised than the laterals, which lie quite at the sides of 
the upper surface of the neck. In F. ocreata the head- and neck- 
stripes are usually badly defined ; when present on the neck they 
are narrow and lie close together. 

In F. sylvestris there is generally a very distinct black wavy 
median spinal stripe, usually extending from behind the shoulders 
to the root of the tail. In /. ocreata the entire spinal area is 
markedly darker than the sides of the body, sometimes showing 
traces of three narrow stripes; but the median is never so strong 
as in J. sylvestris. In the latter, however, a pair of narrow 
interrupted latero-dorsal stripes is sometimes traceable. 

These are the most obvious distinctions. Others are less 
constant. For instance, in /. sylvestris the ears are, generally 
speaking, of the same colour as the head, though not infrequently 
they are washed with yellow either all over or only towards the 
tip. In /. ocreata they are almost always yellower or redder, 
generally very decidedly so, especially in African specimens ; but, 
on the other hand, in an example of /. ocreata sarda in the British 
Museum the yellow on the ear is no more conspicuous than it 1s 
in some examples of /”. sylvestris. 

In F. sylvestris the coat is ionger and thicker than in F. ocreata. 
This imparts to the former Cat a heavier, more robust, and 
shorter-legged appearance, and especially suggests that the tail 
is blanter at the apex *. The value of this difference is discounted 
by the fact that /. sylvestris is a more northern type than 
F. ocreatat, and that the length and density of the fur varies a 
good deal in the latter, which appears to be a species endowed 
with great capacity for environmental adaptation both as regards 
coat and colour. 

Yet, in spite of the obvious resemblances above mentioned, the 
assumption of the total diversity of the two forms seems to have 
been pretty general, if we may judge by the absence of all com- 

arison between them in monographs of the Felide. 

Now the characteristics which the Egyptian and European Wild 
Cats have in common are all possessed by the Domestic Cats of 
the “striped” type; and they are not found in any other species 
of Felis known to me. Hence there is no difficulty in the way of 
believing that our “striped” Cat is the direct and but little modified 
descendant of either /’. sylvestris or Ff’. ocreata, or probably of both 
combined. /. sylvestris inhabits Spain, Italy, Greece, and Asia 
Minor; and /. ocreata Egypt, Tunisia, Algeria, and Sardinia. 
Thus one species or the other is found in the countries bordering 
the Mediterranean basin, where the civilisations of EKurope had 
their origin. If Egyptian Cats were taken to Greece, Italy, or 

* Quite similar differences in the thickness of the tail may be seen in Siberian 
and Indian Tigers. 

+ A very good description and an excellent figure of this species may be found 
under the name Felis lybica in Anderson’s and de Winton’s ‘ Mammals of Egypt,’ 
pp. 171-176, pl. xxiv. (1902). 


Spain, it is highly probable that they interbred with the native 
Wild Cat, especially at a time when the latter was far more 
abundant than it is now and when precautions to prevent the 
tame animals from straying are not likely to have been rigidly 
observed. But even if such crossing took place, I do not believe 
its effects could be traced in the progeny with any certainty, on 
account of the resemblances between the parent forms. 

It may however, in my opinion, be assumed that the differences 
our English striped Domestic Cats exhibit from /’. sylvestris on 
the one hand and from /’. ocreata on the other, coupled with an 
unmistakable likeness to both, are attributable to that cause. To 
assume that the striped pattern of this Cat is due to interbreeding 
between domestic “ Tabbies” or Blotched Cats and specimens of 
F, sylvestris, needlessly complicates a quite simple question. 

It is difficult to decide which of the two species our Striped 
Cats most resemble. Typical short-haired individuals recall 
fF. ocreata in the length of the fur on the body and tail; whereas 
in the colour of the ears greater similarity is presented to 
F. sylvestris than at all events to African examples of /’, ocreata. 
In the proximity of the stripes on the neck, resemblance is evinced 
to F. ocreata ; but their distinctness recalls, though in a lesser 
degree, those of /. sylvestris. In the distinctness of the spinal 
stripe, the domestic form les nearly midway between the two; 
but the transverse stripes on the body and tail are as a rule more 
sharply defined than in typical members of either of the wild 
species: but their want of definition in some long-haired specimens 
is quite paralleled by that of /. sylvestris, and suggests that their 
indistinctness in the latter is attributable to length of fur. 

In the length of the tail the Striped Cat seems to be nearer 
F, sylvestris. There is no evidence that this organ is shorter in 
the domestic form than in the wild species just mentioned ; 
whereas in North-African examples of /. ocreata the tail is 
longer than in either of those types*. 

In connection with the likeness this Cat presents both to Felis 
ocreata and F’, sylvestris, it is apposite to note that out of four 
authors who have described specimens under the belief that they 
represented wild species or races, two compared them to F’. sylvestris 
and two to #. ocreata. Blyth tT, speaking of an example from the 
Punjab Salt Range, says “itis of the streaked or spotted type, 
the colouring and markings of which are not much unlike those 
of the European Wild Cat (/. sylvestris)” ; and Mivart {, in his 
description of /. torquata, remarked, “this Cat has much 
resemblance to the European Wild Cat.” Blanford, on the 
contrary, says “the characters of the upper premolars distinguish 
F. torquata from the allied /. caffra (or caligata) | =ocreata]” § ; 

* Anderson and de Winton, ‘Mammals of Egypt,’ P- 172. 

+ J. A. S. Beng. xxv. p. 442 (1856). t ‘The Cat,’ p. 420 (1881). 

§ ‘Mammals of British India,’ p. 86 (1888). The character Blanford refers to is the 
nearness of the first maxillary premolar to the second—a very variable feature in the 
skulls of Domestic Cats and also, though in a lesser degree, in those of F’. ocreata. 

160 MR. R. I. POCOCK ON [ Feb. 19, 

and Miss Bate described a specimen from Crete as the type of a 
distinct subspecies of /. ocreata *. 

The origin of F’. catws appears to be at present quite unknown. 
Tt may be held :— 

a. That it arose per saltwm as a sport from the other Domestic 
Cat (torquata), and that the pattern persisted im virtue of 
its own inherent dominance without the aid of Man, or in 
virtue of the guiding factor of selective breeding. In 
opposition to this must be urged the complete absence 
of evidence that species of elis are ever dimorphic in 
pattern, and the ascertained fact that they breed true to 
the specific or subspecific type. 

b. That it arose from torqguata by the slow and gradual process 
of preserving and breeding from fancied varieties. But in 
answer to this it may be pointed out that there is no reason 
to think that selective breeding of Cats was ever seriously 
practised until the latter portion of the nineteenth century. 
Moreover, if the catus type arose by that process, inter- 
mediates between it and torguata would probably be seen 

c. That it resulted from the interbreeding of /’. ocreata and 
F. sylvestris. When two distinct species cross, the hybrid 
sometimes reverts in some respects to the characters of a 
common ancestor of both. There is no reason, however, 
for thinking that the pattern of catus was the pattern of 
the ancestor of sylvestris and ocreata; and it seems to be 
in the highest degree improbable that the progeny of two 
closely allied and similarly striped species like sylvestris 
and ocreata should be marked in a totally different manner 
from its parents. There is, moreover, good reason for 
thinking that the torqguata-breed was the resulting hybrid 
of that cross. 

d. That it resulted from interbreeding between sylvestris or 
ocreata or torquata and some exotic species introduced into 
Europe. ‘There is, however, no reason to believe that either 
tamed or wild representatives of any exotic species other 
than ocreata were so introduced, apart from menagerie- kept 

e. That it is the direct descendant of some existing exotic species. 
It is quite evident, however, that it is not the direct un- 
modified descendant of any known species of Felis, since its 
pattern is unique in just the same way and to the same 
extent that the pattern of the Tiger is unique in the genus. 

f. That it is the survivor of some extinct, probably Pleistocene 
Cat of Western Europe. By the method of exhaustion of 
other possibilities, one falls back upon this supposition, 
which at least has this in its favour, that no very obvious 
or cogent reason can be advanced against it. 

* P. ZS. 1906, p. 317. 


4. On Manz, Persian, Siamese, Indian, and Abyssinian Breeds. 

As already explained, ‘‘ Manx” Cats may be either of the catus 
or torquata type. Apart from the abbreviated tail, which has 
been discussed and dismissed as of no systematic significance from 
the zoological standpoint, Manx Cats differ or are alleged to differ 
from ordinary short-haired Cats in standing relatively higher at 
the hind-quarters. I have not seen any measurements sub- 
stantiating this claim ; and it is dificult to decide to what extent 
the greater apparent posterior stature is an optical illusion caused 
by the absence of the tail. It is quite possible, however, that 
suppression of the tail is correlated with greater height at the 
sacrum, and that, to put it crudely, the caudal material is dis- 
tributed to or absorbed by the hind-quarters. Some Lynxes 
certainly seem to stand higher on the hind legs than the majority 
of species of Felis. Be thisasit may, the circumstance that height 
at the posterior region is considered by fanciers ‘“‘ a point” in the 
Manx breed throws the fact—if fact it be—under suspicion of 
having been fostered by selective breeding and of being therefore 
unworthy of consideration from the phylogenetic and systematic 
point of view, zoologically speaking. In other words, as little 
importance should be attached to the character as to the absence 
or abbreviation of the tail. . 

With regard to so-called ‘“‘ Persian ” or ‘* Angora” Cats*, there 
seems to me to be no reason to suppose that any other species is 
involved in their ancestry than in the ancestry of the short- 
haired breeds most common in Europe. Both the “blotched” 
and the ‘striped ” styles of pattern occur; and no other type of 
pattern is known amongst them, so far as I am aware. The 
skull, moreover, is not distinguishable from that of short-haired 
Domestic Cats. The small systematic value that should be 
attached to the long coat has been already insisted upon. Nothing 
seems more likely than that tame Cats were imported from 
Egypt into Persia; the European Wild Cat (7. sylvestris) occurs _ 
both in Asia Minor and Persia, and F. ocreata has been recorded 
from Syria and Arabia. Hence it seems needless to look beyond 
the two species just mentioned for the origin of “striped ” 
** Persian” Cats, even on the supposition that they came ori- 
ginally from Persia. That the ‘‘ blotched ” Persians had the same 
origin, whatever that may have been, as the“ blotched” short-haired 
Cats is in the highest degree probable. For myself, I think 
it quite needless to consider that so trivial a character as long 
hair is to be traced in all cases to Cats imported into Western 
Europe from Asia Minor or Persia. The suggestion—first made, 
I believe, by Pallas, but repeated even in modern literature on 
the subject—that the Central Asiatic species, Pallas’s Cat (Felis 
manul), contributed to the “ Persian” breed, has nothing to be said 
in its favour. Felis manul is quite unlike all domestic breeds. 

* Desmarest (Nouv. Dict. vi. p. 122, 1816) gives Anatolia as the original country 
of this breed. 

Proc. Zoot. Soc.—1907, No. XI. 1] 

162 MR. R. I. POCOCK ON [ Feb. 19, 

The ears are small and set very low on the sides of the head, 
leaving a great width of forehead between them; the facial and 
body markings are different; and, lastly, the pupil of the eye 
contracts to a circular disk *. 

There is, however, one species of Felis which must not be 
altogether forgotten in considering the possible origin of Persian 
Cats. This is the Bokhara Steppe Cat (/elis caudata), described 
by J. E. Gray (P. Z.8. 1874, pp. 31-32, pls. vi. & vii.). The type 
is in the British Museum. ‘The figures of the entire animal and 
of the skull show that this Cat is closely related to Felis sylvestris 
and to /. ocreata, and not to /. chaus, with which Dr. Gray com- 
pared it. It has a thick coat and bushy tail, like 2. sylvestris ; 
but there are no distinct spinal stripe nor definite stripes on the 
head and neck. In length the tail resembles that of /. ocreata ; 
and likeness to this species is further shown by the indistinctness 
of the head, neck, and dorsal stripes. From both the other species 
it differs in being covered with small spots; but these spots, at 
least on the anterior part of the body, show, I think, signs of the 
coalescence into transverse rows which is realised in /’. sylvestris 
and /. ocreata. Gray describes the colour as yellowish. The 
skin, which is probably faded, might be more aptly described as 
‘““ounce”-grey. Whether or not this species has any connection 
with the “spotted” Cats of the Punjab Salt Range, mentioned 
below (p. 164), I am unable to say. 

It is worth putting on record the fact that in two out of the 
three skulls of “ Persian Cats” I possess the jaws are slightly 
“underhung,” that is to say, the mandible protrudes a little in 
front of the premaxillz. I have not noticed this peculiarity im the 
skulls of any other Cats, either wild or domestic. It may be purely 
accidental, or it may indicate that the taste of fanciers in Cats is 
running along the same lines as those of breeders of Domestic Dogs. 
In this case we may in the future have a race of snub-nosed Cats 
departing in facial elegance from Nature’s type of Felis in the 
same manner that Pugs and Bulldogs depart from Nature’s type 
of Canis. 

The origin of the Siamese breed has been a much-discussed 
puzzle. The peculiar coloration fT must be set on one side as value- 
less towards affording a clue. The cats are obviously albescent, as 
is attested by the hue of the hair and frequently by the blueness 
of the eyet. But the albescence, complete in the newly-born 
kitten, is mitigated as age advances by a melanistic taint which 
evinces itself in two ways, both of unusual occurrence in Nature. 
In the first place, the black pigment is distributed symmetrically, 
and primarily affects the face and ears, the tail, and the legs. 

* The illustration of this species in Elliot’s ‘ Monograph of the Felid,’ pl. x., was 
taken from a stuffed specimen, and does not correctly depict the peculiarities of the 
head and eyes. Elliot, however, very rightly insists that this Cat is unlike all other 
known species. 

+ Mr. E. G. B. Meade Waldo has reminded me that the so-called Himalayan breed 
of domestic rabbits is almost identically coloured. 

+ R. I. Pocock, Ann. Mag. Nat. Hist. (7) xix. p. 194, 1907. 


In the second place, it increases in quantity as age advances, and 
gradually encroaches upon the whiter areas, converting the hair 
of the body from whitish or pale fawn into pale or dark brown. 
Sometimes there are indistinct traces of spots on the hind-quarters 
and legs. 

Judging from their size, form, and general aspect, I should say 
these Cats are nothing but a domestic variety of /. ocreata. The 
only skull of this breed that I possess * is that of a female with its 
contours and ridges indicating considerable muscular development. 
It is a short, broad skull, with expanded zygomata, and beyond all 
possibility of doubt falls into the group typified by /. ocreata and 
F’. sylvestris. In profile it is noticeable that the highest point of 
the cranium lies well behind the fronto-parietal suture, and that 
the area in front of that point, almost as far as the nasals, is nearly 
flat and slightly inclined. This is not a common feature in the 
skulls of Domestic Cats; but it is almost exactly paralleled in the 
skull of an old male specimen of /. screata from Suakin, and does 
not occur in the skull of a Siamese Cat in the British Museum, 
in which the highest point of the skull is on the frontal bones, 
which are evidently swolien just behind the postorbital processes. 
Another peculiarity in my specimen is the division of the infra- 
orbital foramen by a bridge of bone. I find this feature repeated, 
however, in the skull of a London Cat not referable to the Siamese 
breed ; and it does not occur in the Siamese skull in the British 

Nevertheless, these two Siamese skulls agree in the possession 
of two small characters, one of which I can only match in one of 
the English Cats’ skulls I possess, while the other cannot be quite 
matched in any of them. The first character is the height of the 
interparietal crest, which is better developed than is usually the 
case in English Domestic Cats, though it is equalled in the skull 
of one London Cat that I have seen, and of one feral example of 
the torquata-type of F. ocreata, from Celebes, in the British 
Museum. The second character is the greater narrowness of the 
posterior portion of the nasal bones and the more marked abrupt- 
ness of the constriction between this portion and the expanded 
anterior portion. But since the two Siamese skulls differ in 
degree with respect to this character, and since the nasal bones 
vary greatly in length and width in English Domestic Cats, no great 
importance can, I think, be attached to the feature in question. 
Moreover, as already stated (p. 152), the Celebes example of 
torquata in the British Museum also has the nasals compressed. 

Since, therefore, the colour of the Siamese Cat affords no 
evidence of its descent, and the skull is decidedly of the ocreata+ 
sylvestris type, there seems to me to be no reason to look beyond 
those species or indeed beyond ocreata for its origin. Those who 
claim that it has had an origin independent of our own Domestic 

* Tam indebted to Mr. F. W. Cousens, F.Z.S., for the skin and skull of one of 
the two specimens of this breed I have been able to examine. Mr. Cousens also 
kindly gave me the skins and skulls of two Persian Cats. 


164 MR. R. I. POCOCK ON [ Feb. 19, 

Cats from some Siamese or Oriental species, must be challenged 
to produce that species before the question can be profitably 
discussed. There seems to be no reason whatever for entertaining 
Trouessart’s suggestion* that the rare Bornean Felis badia was 
its agriotype. I may add that all the small species of Felis in- 
habiting Siam, including even /. chaus, which is doubtfully 
indigenous, differ in the structure of the skull and teeth from 
the Siamese Domestic Cat 7. 

According to Blyth (Journ. As. Soc. Bengal, xxv. pp. 442-445, 
1856) two types of Domestic Cat were prevalent in India in his 
time. ‘One is the streaked or spotted type, the colouring and 
markings of which are not much unlike those of the Huropean 
Wild Cat (Ff. sylvestris), only more distinct, and the transverse 
streaks are more broken into spots, especially towards the hinder 
part of the body ; the fur, however, is short, and the tail £ slender 
and of uniform apparent thickness to the end, showing a series of 
rings and a black tip; ears slightly rufescent externally but 
infuscated, but passing to black at tip, where there is a distinct 
small pencil-tuft of black hairs. Paws deep sooty black under- 
neath.” Two examples of this Cat were shot wild in the Punjab 
Salt Range; another seen at Allahabad in a state of domestica- 
tion exactly resembled them ; but the tame Cats of Calcutta were 
usually greyer, with smaller and more numerous spots. 

The other type, says Blyth, resembles the Jungle-Cat (/. chaus) 
in colour, the body being uniformly “ cat-grey,” more or less rusty 
or fulvescent, without a trace of spots or stripes, but the stripes 
on the tail and limbs are more distinct; there are also confused 
stripes on the forehead and two on the cheeks and a dark band 
across the chest; the lower parts are whitish or tinged with 
tawny, and spotted; the ears are dull rufous behind with a 
slight blackish tip and no pencil of hairs; the paws more or less 
sooty beneath. In its proportions, however, this Cat is quite 
different from /’. chaus, the limbs and ears being much shorter, 
and the tail much longer and tapering. Blyth adds that Domestic 
Cats of this type abounded in Bengal, if not all over India, but 
were quite unknown in Europe; whereas, on the other hand, 
the English “tabby” was quite unknown in India. 

Mr. W. L. Sclater also discussed these two breeds, and accredited 
Blyth with the belief that the self-coloured ‘“ chaus”-like type was 
derived from interbreeding between the ‘‘ Domestic” Cat and 
F. chaus (Cat. Mamm. Ind. Mus. 11. p. 233, 1891). He also sug- 
gested that the form named /. torquata by F. Cuvier was based 
upon a feral example of the spotted or streaked breed. 

So far as I can judge from what the two authors quoted say 
about these breeds, there is nothing to distinguish them from 

* Cat. Mamm. 1904, p. 273. 

+ For list of the species, see S. S. Flower, P. Z.S. 1900, pp. 322-327. 

~ In this particular the tail must resemble and not differ from that typical of 
F. sylvestris. Surely Blyth intended to say “tapering” towards the end, otherwise 
the remark is pointless. 


F’. ocreata, which may be either self-coloured like the ‘ chaus”- 
like Cat or spotted and streaked like the spotted Indian Cat. The 
facts adduced do not appear to me to supply any evidence that such 
indigenous Indian Cats as F’. rubiginosa, PF. chaus, or F’. benga- 
lensis have contributed tothe strains; and I strongly suspect that 
they have been derived from /, ocreata either by the importation 
of tamed specimens or by the reclaiming from the wild state of 
examples of this species which may have inhabited India in com- 
paratively recent times. 

In the British Museum there are a few skins of Indian 
Domestic or semi-domestic Cats belonging to the spotted or 
striped and to the self-coloured types mentioned by Blyth. An 
examination of them fully confirms the opinion I had formed 
from reading Blyth’s remarks. Those belonging to the striped or 
spotted type are, as Mr. Sclater suggested, the same as the form 
described by Cuvier as torguata. ‘Those of the self-coloured type 
do not differ from them more than some of the self-coloured Cats 
differ from the striped Cats to be seen in the London streets. 
Some of them are more rufescent than others; but I cannot find 
in them a particle of evidence of partial descent from 7. chaus 
or from any other indigenous Indian species. 

Not uncommonly Indian Domestic Cats differ from typical 
English examples of torquata in having the bands on the tail 
narrower and the stripes, especially on the forehead and cheeks, 
more rufous. The narrowness of the caudal stripes arises from 
the splitting of the normal stripes; but this character as well as 
the rufescence may be seen in African examples of /. ocreata. 
There is no reason, therefore, to doubt that the Indian Domestic 
Cats are descended from that species. It must, of course, be con- 
ceded that they may interbreed with indigenous Indian species, 
and especially with the so-called Desert-Cat (2. ornata), which, 
from the structure of the skull and teeth, as well as from other 
characters, must be regarded, I think, as the Indian representa- 
tive, as /”. caudata is the Bokhara representative, of the group 
typified by /. ocreata in Africa, and by F’. sylvestris in Kurope. 

Cats of the so-called “ Abyssinian” breed may be descended, for 
anything I know to the contrary, from specimens of 7. ocreata 
directly exported from Abyssinia. They are certainly not unlike 
some self-coloured examples of that species. On the other hand, 
it would I imagine be difficult to separate them from fulvescent 
“ Ticked” Cats, which appear to me to be nothing but examples 
of the torquata-type in which the pattern is broken up and 
evanescent (see p. 152). 

On alleged Cases of Interbreeding between Domestic Cats and 
various wild Species of the Genus Felis. 

It has been stated over and over again that Domestic Cats 
interbreed freely with the native Cats of various species inhabiting 
the countries to which they have been transported. One cannot 

166 MR. R. I. POCOCK ON [ Feb. 19, 

help wishing there was more positive evidence of the fact. It 
may be so; but the statements of authors on this subject cannot 
be accepted as of any great value unless there is collateral evidence 
of their intimate acquaintance with the wild species in question 
and with the range of variation in colour, pattern, and structure 
of the domestic breeds. Many alleged cases of interbreeding may 
be found quoted in Darwin’s ‘ Variation of Animals and Plants 
under Domestication.’ It is very likely that some Cats seen 
by Jardine in the north of Scotland were rightly regarded as 
hybrids between domestic animals and J. sylvestris. And 
St. Hilaire’s statement that the Domestic Cats of Algiers cross 
with the native Cat (/. lybica = ocreata), and Layard’s to the 
effect that similar crosses occur with /. ocreata caffra in 
S. Africa, may also be true. But in the last two cases it may be 
doubted if evidence of the crossing would be shown by the progeny ; 
and as regards Jardine’s alleged hybrids, long-coated specimens of 
the “striped” Domestic Cat might easily be mistaken for half- 
bred Wild Cats. A striped Cat known to be descended from a 
feral specimen was recently caught in the New Forest, where 
Felis sylvestris has not occurred for at least a century, and sent to 
the British Museum by Mr. P. H. Barker as a hybrid. On the 
strength of its long coat, it must be regarded as a ‘“ half Persian.” 
Had it come from parts of the north of Scotland, where /. sylvestris 
still lingers, the entire purity of its descent might have remained 
for ever in doubt. As regards the claim that Domestic Cats in 
India interbreed with the Jungle-Cat (/’. chaus), I am unable to 
find evidence that satisfies me of the certainty of this occurrence 
(cf. supra, p. 165). In one case the claim is actually based, in 
part at all events, upon the occurrence of the internal brachial 
stripe on the fore leg, a feature regarded by the observer as 
characteristic of F’. chaws, whereas it is the most persistent of all 
the stripes in the genus els. 

No one would be so rash as to atlirm that interbreeding does 
not occur between Domestic Cats and even widely different wild 
species. But unless the skins and skulls of alleged hybrids are 
forthcoming for examination, there is no basis for the discussion 
of the question. Vague suppositions of observers cannot be 
regarded as evidence of the fact; and I am of opinion that the 
prevalent beliefs on the subject are to be assigned in a great 
measure to the observation of semi-wild specimens of the common 
but little known striped Domestic Cat, which may be either red 
or grey in colour, and either striped or spotted in pattern. 

6. Summary. 

The substance of the foregoing remarks may be epitomised as 
follows :— 

1. The characters used by breeders and fanciers as a basis for 
their so-called breeds of English Domestic Cats have no 
scientific value, in the sense of affording a clue to affinity 
and descent. 



. The pattern—or, in other words, the arrangement of the 
stripes—shows that English Domestic Cats are referable 
to two distinct types, whether they belong to the ‘‘ Manx,” 
‘“« Persian,” or ‘ Short-haired ” breeds. 

3. These two types of pattern are different in kind and do not 
intergrade. They are so distinct from each other that no 
one would hesitate to regard them as characterising two 
well-marked species if the animals presenting them existed 
in a wild as opposed to a domesticated state. 

4, In one type of pattern the stripes take the form of narrow 
transverse or vertical bands which sometimes break up 
into spots. To feral or domesticated examples of this Cat 
have been given many names, of which torquata is the 
best known and angorensis or striata possibly the oldest. 

5. This Cat (torquata) was apparently domesticated in Europe 
at least as early as the 16th century. There seems to be 
no reason therefore for regarding it as of Indian origin. 

6. It closely resembles in pattern two existing species, namely, 
the so-called Egyptian Cat (/. ocreata) and the European 
Wild Cat (/. sylvestris), both of which occur at the 
present day in the Mediterranean Region, and are very 
nearly related to each other. There is no difficulty in 
the way of believing that they are the ancestral forms or 
‘“‘aoriotypes” of this domesticated race (torquata). 

7. In the other type of pattern the stripes take the form of 
broad longitudinal or obliquely longitudinal bands forming 
a ring-like or spiral arrangement on the sides of the abdo- 
men. To domesticated examples of this Cat, Linneus gave 
the name catus, which cannot be applied to any other form 
of the genus Felis. Domestica is its best-known synonym. 

8. This Cat (catus) is certainly known to have been domesti- 
cated in Europe in the middle of the 18th century. It 
was not, however, apparently known in India in the 
middle of the 19th century. Probably, therefore, it is of 
European descent. 

9. Its origin is unknown. Of the several hypotheses that may 

be held on this subject perhaps the following two are the 

most to be commended :—that it arose as a sudden varia- 
tion or sport from the forquata-breed, in which case 

European Domestic Cats are dimorphic in pattern; that 16 

is the direct descendant of some extinct Pleistocene Cat, 

in which case there are two distinct species of Domestic 

Cat in Europe. 


Puate VIII. 

Blotched Tabbies, Felis catus. 

The four skins photographed were selected out of a large number to show the 
principal variations to which the pattern in this Cat is liable. All came from the 
Cats’ Home in Camden Town, London. 

168 DR. C. G. SELIGMANN ON DEATHS [Feb. 19, 

Prats IX. 
. Striped Tabby, Felis torquata. 

Fig. 1. Example of the Felis sylvestris-type to be compared with fig. 1, Plate X. 

2. Like the last, but the pattern more spotted. 

3. A partially albino specimen with the spots larger and more widely spaced 
than in No. 2. 

4. Specimen of the so-called “Ticked ” breed, with the spots disintegrated and 
generally distributed over the body. 

(From the Cats’ Home, Camden Town.) 


Puate X. 
Agriotypes of the Striped Tabby, 7’. torquata. 

Figs. 1 & 2. European Wild Cats, F. sylvestris, from Ross-shire in Scotland. 

Figs. 3, 4, 5. Young examples of the South African race of the African Wild Cat, 
Felis ocreata caffra. The pattern is usually more distinct in the young 
than in the adults of this species. The exact lscality of these three 
specimens is unknown. They were shipped from Cape Colony. 

(The camera has emphasised the pattern of the skins depicted on this Plate.) 

Report on Deaths occurring in the Society’s Menagerie 
during 1906. By C. G. Seniemany, M.D., F.Z.8., 
Pathologist to the Society. 

[Received February 19, 1907. | 
(Text-figure 61.) 

In the annexed table will be found the causes of death so far as 
they could be discovered of 355 mammals and 283 birds which 
died in the Society’s Gardens, and which were submitted to post- 
mortem examination during the year 1906. In these mammals 
and birds no cause of death is stated to have been found in 36 
mammals and only one bird, but 1t must be noted that the number 
of deaths put down to trauma and exhaustion is much larger than 
in 1905. This is due, in my opinion, to the important part 
played by depressed vitality, and in some cases darkness and 
cold, in bringing about death. The method of classification is the 
same as that adopted last year*, but two new headings occur, viz. 
Diseases of the Ductless Glands and Deaths due to Old Age. 
Possibly two of the deaths among birds attributed to “ trauma 
and exhaustion” really took place owing to the effect of parasites. 

The following remarks refer to conditions of special pathological 
interest occurring in the animals in the Gardens. 

Tuberculosis. ae is yet too early to attempt any full appreciation 
of the effect of thoroughly disinfecting, scraping, and repainting 
the monkey-house, undertaken early in the year, when the heating 
arrangements also were altered, but the diminution of deaths from 
tuberculosis—34 in 1906 against 53 in 1905—is decidedly 
encouraging. If the figures be expressed as percentages, 21°6 of 
the total deaths in monkeys were from tuberculosis in 1906 against 

* “Note on Deaths occurring in the Society’s Gardens during 1905,” P. Z.S. 1906 
p. 234 et seq. 




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170 DR. C. G. SELIGMANN ON DEATHS [ Feb. 19, 

35°8 in 1905. Assuming that there were about 100 monkeys in 
the house on January the Ist, 1906, between 16 and 17 per cent. 
of the inmates of the house have died of tuberculosis during the 
past year, since the actual number of monkeys received was 105. 

My thanks are due to Mr. Pocock for ascertaming the number 
of admissions during the year, and I am further indebted to him 
for the estimate of 100 as the number of monkeys in the house 
in January 1906. 

Much attention has been paid to tuberculosis in birds, and 
certain interesting conclusions concerning avian tuberculosis can 
be drawn from a study of the material available. It must in the 
first place be noted that during the past two years about 30 per 
cent. of all deaths in birds have been due to tuberculosis; and the 
more the matter is Investigated the more obvious it becomes that 
a very large percentage of the cases of tuberculosis occurring in 
birds in the Zoological Gardens are the result of infection by the 
intestine, which can hardly be due to any other cause than the 
swallowing of particles of contaminated soil while food is being 
picked up. By far the greater proportion of birds dying of 
tuberculosis in the Gardens present typical lesions in their spleen 
and liver, which can only be explained on the hypothesis of an 
ingestion tuberculosis: sometimes the intestine is affected, but 
more often this is not the case, and typical tubercular ulceration 
of the gut in birds dying in the Gardens is rare. But although 
ulceration of the gut does not frequently take place, enlarged 
tuberculous glands at the root of the mesentery are by no means 
uncommon; a condition akin to tabes mesenterica of human 
pathology being set up without obvious damage to the mucous 
membrane of the gut, but with the addition of lesions in the 
spleen or liver or both. Even where the intestine is affected, 
ulceration is rare or slight; comparatively large submucosal 
nodules being formed over which the villi of the mucous mem- 
brane often seem enlarged, so that a curious condition suggestive 
of multiple closely-set warts is found to occupy the inner surface of 
the bowel. Sometimes, as in a Vulturine Guinea-fowl (Acryllium 
vulturinum), the whole of the large gut may be thickly studded 
with these warty growths, which on section are found to 
contain dense masses of acid-fast bacilli. In other cases the 
whole of the small gut is similarly affected, as in a Burmese Slaty- 
headed Parakeet (Palecornis sp.). In the latter case it was inter- 
esting to note that there were no lesions in any other abdominal 
organ or in the thorax, while in the case of the Guinea-fowl there 
was early tuberculosis of the lung, liver, and spleen. 

Pneumonia and Broncho-pneumonia._—No marked improvement 
in these diseases has followed the cleaning of the monkey-house. 

Mycosis.— Reference was made last year to the occurrence of a 
disease, mycosis, which was due to the invasion of the tissues by 
a mould, Aspergillus fumigatus. This disease is by far commoner 
in water-fowl than in other birds, and when attacking these its 
characteristic lesions are usually widely distributed throughout 
the body-cavity ; but a case occurred in an African Buzzard (Buteo 


desertorum) in which the disease, obviously not very recent, affected 
only the air-sac and the lung of the right side, there having been 
no extension of the parasite by continuity or by infection of the 
blood or lymph stream. 

Cardiac Failure —It has been noted in the case of Ostriches, 
Rheas, and Cassowaries, as well as some of the larger Storks kept 

Text-fig. 61. 

Aorta of Tiger, showing several aneurysms. 

in the Gardens, that their hearts after death are usually extremely 
flabby, while the subpericardial fat may be replaced by a loose- 

172 DR. C. G. SELIGMANN ON DEATHS [ Feb. 19, 

meshed cedematous tissue; so that probably cardiac failure is the 
direct cause of death of many of these birds, the condition perhaps 
being due to lack of exercise. One case of particular interest as 
bearing out this view has occurred during the past year. While 
some Zebras were being moved into the new enclosure in the 
neighbourhood of the Seal pond,an Emu (Dromews nove-hollandie), 
so placed as to be able to see what was happening, became 
extremely excited, and running round its paddock either struck the 
railings and collapsed against anes or else collapsed and fell against 
the railings. In any case it could hardly have been injured by the 
blow, since its feathers were not damaged or its skin torn. On 
picking it up it was found to be quite dead, but at the post-mortem 
examination no sign of disease could be found in the brain, 
or abdominal organs, nor was the heart notably flabby. 

Arterial Disease-—The aorta showing many aneurysms of an 
old Tigress, which had lived in the Gardens for 1323 years, was 
shown at one of the evening meetings of the Society, and a brief 
note upon the condition appeared in the ‘ Proceedings’ *. An 
illustration has since been prepared which is reproduced in text- 
fig. 61, and shows the unusual condition of the vessel, the previous 
description of which may be quoted here :—‘ The aorta shows 
advanced arterial disease, most pronounced in the descending 
aorta, where there is marked atheroma and where, in a length of 
about 180 mm., there are 14 aneurysmal swellings varying in size 
from that of a pea to that of a fair-sized plum. The two largest 
swellings, the walls of which are of stony hardness, occur close 
together on opposite sides of the artery.” Arterial disease, though 
not common in the animals in confinement in the Gardens, cannot 
be said to be rare, whereas in wild animals this disease is generally 
considered to be very.rare. Perhaps an interesting parallel may 
be drawn in this respect between man under civilised conditions 
and animals in confinement. In the former arterial disease is of 
course common, but there is considerable evidence against its 
occurrence among certain people but just emerging from the Stone 

Gastric Ulcer.—The experience of this and last year seems to 
show that if gastric ulcers are not entirely confined to the 
Carnivora and Marsupialia, they are at least most common in 
these orders, since of a total of nine cases four occurred in 
Carnivora and five in Marsupialia. A young Ocelot (felis par- 
dalis) presented the lesions of this disease in a_ particularly 
interesting form. On opening the belly, part of the colon looked 
darker than usual, and it was found that this and the rectum 
were full of dark semi-digested blood; there were many small 
recent hemorrhages in the stomach, over some of these the mucosa 
was destroyed but the ulcers had not penetrated deeply. In the 
duodenum there were, however, sixteen orifices, all more or less 
circular, and varying In size from that of the head of a large pin 
to that of a threepenny-bit. One of the largest of these extended 

* P.Z.S. 1906, p. 634. 



as deeply as the serous coat, the others were less deep and 
involved only the mucosa and a part of the muscular walls of the gut. 
The Marsupials with gastric ulcer were all herbivorous. 

Perforation of the Uterus.—An instance of this unusual accident 
oceurred in a bitch of the North-African Jackal (Canis anthus). 
The mammary glands betokened recent activity, and on opening 
the abdomen the uterus projected well above the pelvis; there was 
general peritonitis and the uterus itself was dark and intensely 
congested. Within it lay a dead and extremely offensive fcetus, 
while a small circular perforation existed in the uterine wall just 
above the cervix on the left side, and it was apparently due to 
leakage through this hole that peritonitis had set in. 

New Growths.—The rarity of new growths referred to in last 
year’s note is confirmed by this year’s post-mortem examinations 
on a larger number of animals. Only four cases occurred ; two of 
these, both in mammals, were carcinomata and presented no 
unusual features; but the third case, a Bear, had multiple small 
angeiomata of the liver, none bigger than a sixpenny-bit, which 
cannot have exerted any evil effect on the health of the animal. 

The last case occurred in a Chilian Pintail (Dafila spinicauda) 
alleged to have been bred in the menagerie and to be 26 years old. 
On opening the bird a mass the size of a turkey’s egg was seen 
occupying the right flank in the front of the belly; it was not 
adherent to the intestine or other organs but was enclosed within 
a thin, loose capsule resembling peritoneum within which, with 
the mass, were the supra-renals, which were not affected. The 
mass could not be traced to any organ, but seemed to arise at the root 
of the mesentery; no testes could be found, although the trachea 
was of the normal male type; there was a white mass about the 
size of a filbert in the right lobe of the liver, and both kidneys 
were whitish and much enlarged. Microscopically it was seen that 
the main mass was structurally a carcinoma, as was that in the liver, 
and both these masses resembled the much enlarged kidneys which 
were themselves carcinomatous. 

My thanks are due to Mr. 8. G. Shattock for the diagnosis 
of this remarkable case, which it seems must be classed as an 
example of diffuse carcinomatous growth in both kidneys with 
secondary masses in the liver and glands at the root of the 

While considering the occurrence of new growths in birds, 
allusion may be made to two very interesting conditions which 
have recently come under my notice. The subject of the first of 
these was a domestic fowl belonging to Dr. R. N. Salaman * 
which died suddenly. On opening the belly cavity this was found 
to be full of blood, and the liver was found to be diffusely angeio- 
matous. The second condition occurred in a Sparrow examined by 
Dr. A. Wilson * in the course of certain physiological work. On 
opening the skull a spherical mass about the size of a pea was seen 

* My thanks are due also to these gentlemen for allowing me to refer to their 

174 MR. J. T. CUNNINGHAM ON [Feb. 19 

to press upon the right half of the cerebellum, which it had to 
some extent excavated. This mass was enclosed in a sheath of 
pia mater continuous with that over the posterior part of the 
right cerebral hemisphere. Dr. Wilson thought that the spherical 
mass might at one time have been joined to the cerebral hemi- 
sphere, but of this there was no direct evidence. Microscopically 
the tumour mass contains true nerve-cells, and so is an example of 
an extremely rare condition. 

3. On a peculiarly Abnormal Specimen of Turbot. 
By J. T. Cunnincuam, M.A., F.Z:8. 
[Received January 23, 1907. | 
(Plate XI.*) 

The specimen here described was sent tome by Dr. EH. J. Allen, 
Director of the Marine Biological Laboratory, Plymouth, in the 
beginning of December 1906, with a request that I should examine 
and describe it. With the fish was a normal specimen and two 
letters referring to them—one from Mr. John D, Enys, the other 
from Miss Olivia L. Fox. Mr. Enys’ letter is dated Nov. 3, 1906, 
and states that Miss Fox had then alive ina glass globe two 
small Turbot caught on the sands at Polzeth, near the Doom Bar 
at Padstow, on the north coast of Cornwall; that the abnormal 
fish was dark on the under side and white on the upper. Miss Fox’s 
letter states that she had had the fish about a month, and that the 
upper side “‘ was becoming pigmented” since she first obtained it. 

The specimen is 4:4 em. in length and presents a condition 
which, so far as I am aware, has never previously been observed 
or described in flat-fish of any species. With respect to the 
position of the eyes, the fish is a reversed specimen—that is to 
say, both eyes are on the right side, whereas normally in Turbot 
they are on the left. With respect to colour, on the contrary 
the specimen partially resembles a normal Turbot. The right side 
is almost entirely unpigmented ; the greater part of the left side 
is coloured like a normal Turbot. The pigmentation does not 
extend uniformly over the whole of the left side, but is absent 
from the head, and from the anterior part of the dorsal region 
above the head. On these areas there are only a few scattered 
black chromatophores. On the right or uncoloured side there are 
also scattered black chromatophores rather more numerous than 
on the left side of the head. It is important to note that the head 
and anterior region of the right side, although not fully pigmented, 
have more pigment than the rest of that side; between the eyes 
and around the dorsal eye pigmentation is almost complete. 

The number of dorsal fin-rays in the specimen is 65, of the 
ventral 47. The characteristic tubercles of the adult Turbot are 
not yet developed, but there are three little projections at the 
base of each of the dorsal and ventral fin-rays, and also projections 

* For explanation of the Plate, see p. 181. 

Ie eo Wi. Ih. 2: 

d.Greeen hth. 

J.T. C del. 



at the bases of the caudal and ventral rays. These are probably 
the beginnings of marginal tubercles. 

The anterior end of the dorsal fin and the basal tissue which 
carries it forma projecting hook-like process over the dorsal eye— 
that is, the originally left eye, which has moved to the right side 
of the head. This projection, due to the absence of attachment 
between the base of the fin at the anterior end and the head, 
occurs commonly in ambicolorate specimens of the turbot and less 
frequently in ambicolorate specimens of other species of Plewro- 
nectide (see Cunningham and MacMunn, “ Coloration of Skins of 
Fishes, &c.,” Phil. Trans. 1894), 

A letter from Miss Fox to Dr. Allen, dated Jan. 7, 1907, 
explains that the fish was caught on Sept. 28 last year, and 
lived in captivity till Nov. 28, when it died from some unknown 
cause. When caught it was, unlike all the others seen at the same 
time, quite stationary on the sand, which Miss Fox thought might 
imply a certain blindness. In captivity, however, it was very 
active, and certainly saw food very quickly, so that there is no 
reason to think the function of the eyes was affected. 

In Plate XI. I have figured the two sides of the abnormal 
specimen. The normal specimen was 4°2 cm. long. Its meta- 
morphosis is complete, but there are still a few scattered black 
chromatophores on the right or lower side. Similar black 
chromatophores are present on the right side of the abnormal 
specimen, and they appear to be larger and slightly more 
numerous ; but the difference is slight, so that the exposure of 
the right side to light during the two months it was in captivity 
had not produced much effect. 

It seems to me that the only way to attempt an explanation of 
the condition of this specimen is to base the explanation on the 
view of the constitution of the ovum which was developed by 
Weismann, and which is adopted in the Mendelian doctrine of 
heredity. If the right side of the anterior or cephalic region were 
more completely pigmented, we might regard the fish as consisting 
of an anterior smaller part which was reversed, and a posterior 
part which was normal. The condition would then be explained 
by supposing the specimen developed from an ovum consisting of 
parts usually occurring in separate ova. We know that reversed 
specimens occur in various species of flat-fishes, e. g. the Flounder 
(Pleuronectes flesus). In this species in some localities reversed 
specimens are not only common, but abundant. At Plymouth I 
found about 30 per cent. of the specimens captured had the eyes 
and pigment on the left side instead of on the right. It is neces- 
sary here to consider the precise terms to be used to indicate the 
structural peculiarities which present themselves. It has been 
usual to speak of a Flounder with eyes on the left side as reversed ; 
but if we use the substantive corresponding to this adjective, 
namely reversion, we are using a term which has been employed 
in an entirely different sense, namely as meaning atavism, or the 
recurrence in a species of some more or less remote ancestral form. 

176 MR. J. T. CUNNINGHAM ON | Feb. 19, 

It is true we might use the word reversal, but this is not suffi- 
ciently distinct. In order to avoid confusion it will be better to 
coin a new term, and it seems to me the most appropriate term is 
“‘metastrophe,” meaning a change in the direction of the turning. 
For adjectives we may use merely sinistral or dextral, referring to 
left or right side, or for the abnormal condition in general we may 
use metastrophiec. 

Since, then, metastrophe frequently occurs in flat-fishes, and is 
a congenital abnormality due to some abnormality in the constitu- 
tion of the ovum, it is intelligible that it should occur in one part 
of a fish and not in another. We may suppose the abnormality 
in the whole fish is due to the interchange of position in the ovum 
of the parts corresponding to the left and right sides of the body. 
The abnormality does not, however, affect the viscera, which, as I 
have pointed out in the memoir already cited, are constant in 
position whether the fish is dextral or sinistral. In the particular 
specimen of Turbot which we are considering, the head is dextral, 
or metastrophic, the posterior portion normally sinistral, and its 
origin is to be attributed to a corresponding abnormality in the 
constitution of the ovum from which the fish was developed. 

With regard to the question of the origin of such abnormalities 
in the ovum, they may arise either in the cell-divisions which 
occur in the multiplication of ova or spermatozoa of gametes, 
to use the general term, or in the process of fertilisation, the 
conjugation of the gametes. It might be suggested in this 
particular case that the condition was due to a “‘ cross” between 
an abnormal dextral specimen and a normal sinistral specimen, 
the condition of the head-region being inherited from one parent 
and that of the posterior region from another. But metastrophic 
or dextral specimens are, so far as my experience goes, rare in the 
Turbot, and it seems equally possible that the peculiar condition 
of the gamete which gave rise to the abnormality was not due to 
the condition of one of the parents. 

Tt is not necessary to suppose that both of the gametes which 
produced the fertilised ovum were abnormal : abnor mality in one 
only may have been sufficient to produce the abnormality of 
development. In the division of the gametes within the repro- 
ductive organ of a parent fish, the chromosomes of the nucleus, 
which are supposed to be the “carriers of heredity ” or to contain 
the “determinants” which produce the characters of the organism 
to which the gamete gives rise, normally divide severally so 
that two similar ova are produced. In the final or reduction 
division each chromosome does not divide, but the group of 
chromosomes separates into two groups. In one or other of 
these divisions the determinants might be displaced, so that either 
all or some of those belonging to the left side were on the right 
and vice versd, and thus a metastrophic gamete would be produced. 

One important question that arises from the condition observed 
in the specimen under discussion is, what bearing it has on the 
experiments carried out by me some years ago at the Plymouth 


Laboratory, and described in the paper already cited. In those 
experiments pigment was developed on the lower sides of Flounders 
as a result of the incidence of light. Here we have a specimen 
of Turbot in which the upper side is exposed to light and is not 
pigmented, while the lower side is pigmented. But it must be 
noted that no adult specimen has been observed in which this 
condition occurs. According to Miss Fox’s letter quoted above, 
the upper side of this young turbot had already acquired some 
pigment during the two months in which it lived in her possession. 
It is quite possible therefore that if the specimen had lived to 
become adult, the upper or right side would have become fully 
pigmented in consequence of the action of light, and then the 
specimen would have been exactly similar to other ambicolorate 
specimens of Turbot, except that it was metastrophic, the eyes 
being on the right side instead of the left. 

In my experiments, I showed that when young fish in process 
of metamorphosis were placed in the apparatus so that light fell 
on the lower side and not on the upper, the normal hereditary 
changes were not arrested, pigment disappeared from the lower 
side as under normal conditions, and it was only later, after 
long exposure to light, that pigment was developed on the lower 
side. Thus, as the specimen we are here considering had not long 
passed its metamorphosis, there is nothing inconsistent with my 
results in the absence of pigment from the tight side, although that 
side is uppermost and had been exposed to light for a short time. 

The condition of the specimen here described suggests that the 
usual ambicolorate abnormality is due also to partial metastrophe, 
but that in these cases the anterior part of the body is normal or 
sinistral, and the posterior part dextral. This view would explain 
the remarkable fact, of which hitherto no explanation has been 
given, that in the great majority of ambicolorate Turbot the lower 
or right side of the head is unpigmented, just as in the specimen 
here described the left side of the head is unpigmented. The 
limits of the pigmentation are not absolutely constant. In the 
majority of specimens which I have seen, the pigmentation 
extends on to the lower jaw and the anterior end of the dorsal fin, 
while the rest of the head in front of the preopercular bone is 
unpigmented. One specimen in my list, however, had pigmenta- 
tion over the whole of the lower side, including the head. If the 
explanation suggested is correct, it follows that the young of an 
ambicolorate specimen immediately after metamorphosis is without 
pigment on the postcephalic portion of the upper cr left side, and 
that it becomes ambicolorate in adult life in consequence of the 
development of pigment on that side under the influence of light. 
There is at present no direct evidence of this beyond the occurrence 
of the specimen described in this paper, and the question must be 
further investigated by the examination of large numbers of 
young specimens. When pigmentation extends over the whole of 
the lower side, mcluding the head, it cannot be said that the head 
of the fish is normally asymmetrical; therefore the theory of 

Proc. Zoou. Soc.—1907, No. XII. 12 

178 MR. J. 1. CUNNINGHAM ON [Feb. 19, 

partial metastrophe does not apply. In this case we must conclude 
that some other explanation is to be sought, or we may suppose 
that the boundaries between the determinant groups in the ovum 
are not definite, and that the pigment determinants displaced to 
the right side have extended to the head-region. 

It may be objected that the persistence of colour on the lower 
side of an ambicolorate Turbot is inconsistent with my views of 
the action of light, that if pigment were produced on the upper 
side it ought to disappear from the side turned to the ground. 
This objection is of little weight, for my experiments show that it 
is easier by means of light to produce some pigment where it was 
previously absent, than to abolish it when it is present, by cutting 
off the light. This is what might be expected, for in the evolution 
of a flat-fish pigment has only recently disappeared from the lower 
side, in consequence, as I believe, of the absence of light; and 
therefore the pigmentless condition is not very strongly inherited, 
and pigment is produced after a comparatively short exposure to 
light. The positive character on the other hand, the presence 
of pigment, has existed not only since the flat-fish was evolved, 
but in a long line of ancestors before that, and therefore it would 
probably take several generations to cause the pigment to disappear 
completely by cutting off the light. It is quite possible that when 
the lower side is congenitally pigmented, some proportion of the 
pigment is lost in consequence of the absence of light, but such a 
loss would not be obvious to observation and would be difficult to 
demonstrate. Obviously a small amount of pigment appearing 
where there was none before 1s evident at once, but the disappear- 
ance of a small proportion from a strongly pigmented surface 
makes no apparent difference to the colour, and there is no means 
of measuring the amount of pigment for comparison in different 
cases. There can be no doubt concerning the presence of a single 
sheep in a field, but 1t is much more difficult to decide whether 
there are a thousand or 999 in a flock. 

Tt has long been known that in Pleawronectide generally, and 
especially in Rhombus maximus, there is a marked correlation 
between ambicoloration and the malformation of the dorsal fin 
which occurs in the specimen described in this paper. It seems to 
be generally supposed that in such specimens the dislocated eye 
has not completed its change of position, and being on the edge 
of the head instead of on the upper side, prevents the usual growth 
forwards of the base of the dorsal fin. The condition is regarded 
then as, like the ambicoloration, a reversion on the part of the 
eyes and skull towards the primitive symmetry. Although I have 
not fully investigated the structure anatomically, it 1s my opinion, 
from external observation, that the eyes and skull are normal and 
that the peculiarity is merely due to a want of that attachment 
which normally occurs between the base of the fin and the skuil, 
along the united ectethmoid or prefrontal, and frontal bones. T he 
view L have suggested seems to me to give a better explanation 
of this abnormality than has hitherto been proposed. If the head 

1907. ] AN ABNORMAL TURBOT, 179 

is metastrophic and the poster lor region normal, as in the specimen 
here described, or vice versd, aS in ambicolorate specimens pre- 
viously described, then the normal relation of the determinants of 
these parts in the ovum, and therefore in development, is wanting. 
The anterior end of the dorsal fin belongs to the posterior of the 
two portions abnormally joined in the fish. It tends to grow 
forward, but in the normal case in doing so unites with the right 
side of the skull (in the Turbot); whereas in the abnormal specimen 
here described, where the head is metastrophic, it has the left side 
of the skull opposite to it, and with this side it has no congenital 
relations, and so remains separate from it. In the more usual 
case, where the eyes are on the left side as usual but the fish is 
ambicolorate, a similar explanation would apply. Here the right 
side of the skull is opposite the fin, as in the normal fish; but the 
fin being itself metastrophic, the normal relations between fin 
and skull in development are disturbed, and consequently they 
remain separate. It may be said in fact that in all these cases 
the fish. or the ovum from which it develops, is composed of two 
separate parts united in an abnormal relation to one another in a 
plane transverse to the long axis of the fish. Consequently the 
normal continuity between the head and body is, as it were, 
imperfect ; and in all probability this is the real reason why in 
these cases the anterior end of the dorsal fin remains unattached 
to the head. 

The abnormality of the dorsal fin does not occur in specimens 
which are entirely metastrophic. Here, although the characters 
of the right side develop on the left, and vice versa—that is to 
say, the determinants of the right and left sides have changed 
places—the dislocation of determinants in the gamete has taken 
place along the median plane, and therefore the longitudinal 
continuity between fin and skull is not disturbed. 

It is important to mention that the abnormality of the fin in 
the specimen here described is not merely due to incomplete 
metamorphosis. The normal specimen of the same size, or rather 
smaller, sent with the abnormal, and captured at the same time, 
shows complete metamorphosis, and in it the dorsal fin extends 
forward attached to the head to a point anterior to the eyes. 

The correlation between ambicoloration and the abnormality of 
the dorsal fin isnot invariable. Cases occur in which ambicolorate 
specimens are in this respect structurally normal. In the Phil. 
Trans. memoir by myself and Dr. MacMunn (referred to above, 
p- 175), I made the generalisation from the specimens of Turbot 
then known to me, that if pigment was present over the whole of 
the body behind the pre-opercular bone, and also on the lower jaw 
and the anterior end of the dorsal fin, the malformation of the dorsal 
fin was present; whereas if the pigment was less than this, the 
malformation was absent. On the hypothesis of the cause which 
T have suggested, the absence of the malformation inthe latter case 
is intelligible, for then the junction between the metastrophic and 

the normal parts of the body may be supposed to occur not between 

180 ON AN ABNORMAL TURBOY. [ Feb. 19, 

the dorsal fin and the head, but in the course of the dorsal fin 
itself. The anterior end of the fin then belonging to the same part 
of the composite fish as the head, the relations of the two struc- 
tures in development are not disturbed. Holt *, however, records 
aspecimen of Turbot in which there was some pigment on the lower 
side of the jaws and on other parts in front of the preoperculum, 
and yet the head and fin were structurally normal. ‘This excep- 
tion is not enough to disprove my hypothesis, for it may happen 
exceptionally that the two parts in the composite gamete are so 
exactly fitted together in their new relations, that the fin attaches 
itself to the skull in development as in a normal specimen. 

The fact which gives most support to my hypothesis, is that in 
the great majority of ambicolorate Turbot the lower side of the 
head is white and destitute of pigment. The same condition has 
been seen by me in two specimens of Pleuronectes macrocephalus 
and one specimen of Plaice, P. platessa. Inthe Flounder (P. flesus), 
however, I have not found the absence of pigment from the lower 
side of the head in ambicolorate specimens. In the Phil. Trans. 
memoir I have recorded four specimens in which the lower side 
was completely coloured and in which the usual abnormality of 
the dorsal fin was present. 

There is, however, another fact concerning ambicoloration 
which is difficult to reconcile with the view that it is due to the 
metastrophe of the posterior region, namely, that in ambicolorate 
specimens not merely both sides are pigmented, but both sides are 
equally armed—that is to say, the scales, spines, or tubercles are 
equally developed on both sides. In normal flat-fishes the arma- 
ture is much reduced on the lower side. In the Turbot the 
scattered tubercles present on the upper side are almost entirely 
absent from the lower side ; in the Flounder the rough spiny scales 
along the lateral line and along the bases of the dorsal and ventral 
fins are absent from the lower side. In ambicolorate specimens 
the armature is not only present on the lower side, but also on 
the upper; the abnormal specimens are not merely ambicolorate, 
but ambiarmate. If, according to the hypothesis I have suggested, 
the postcephalic region were metastrophic, the upper side should 
be originally not merely without pigment, but without armature, 
like the lower side in normal specimens. If the pigment on the 
upper side in ambicolorate specimens were due to light, the arma- 
ture should remain absent, unless the action of light produces 
armature as well as pigment, for which there seems no reason and 
for which we have no evidence. Something might be attributed 
to the absence of friction from the upper side, but this is excluded 
by the strong development of the armature on the lower side in 
these abnormal specimens. These considerations are in favour 
of the alternative hypothesis to explain ambicoloration and ambi- 
armature, namely, that in the gamete two right or two left sides 
are united instead of aright and left. In this case of course all 

* E. W. L. Holt, “Studies in Teleostean Morphology from the Marine Labora- 
tory at Cleethorpes,” P. Z. S. 1894, p. 413. 


the characters of the upper side would be reproduced in the lower, 
both pigmentation and armature. The symmetry does not extend 
to the eyes and skull, but then the malformation of the head may 
be due to a partial symmetry, a reduction of the normal asymmetry. 

It is possible that both cases occur in different specimens—that 
is to say, that in some ambicolorate specimens the condition is 
due to metastrophe of the posterior region of the body ; in others 
to secondary symmetry, the doubling of the upper side in the 
gamete. This cannot be decided without further investigation of 
abnormal specimens both in the young state and theadult. There 
can be no doubt of the importance of the unique condition 
exhibited by the specimen here described, or that its condition 
is best explained on the view I have suggested, namely, that it 
consists of a metastrophic head joined to a normal body. 

I beg to offer my thanks and congratulations to Miss Fox and 
Mr. Enys for making the specimen known, and my thanks to 
Dr. Allen for allowing me to describe it. 


Abnormal Young Turbot. 


.1. Right (upper) side of abnormal Turbot 4:4 mm. long, enlarged. The eyes 
are on the right side of the head, and the dorsal fin projects anteriorly as 
a free process. Some pigment on the head dorsally, elsewhere only 
scattered black chromatophores. 

g. 2. Left (lower) side of the abnormal Turbot shown in fig. 1. No eyes on 

the lett side of head, nor pigment. Pigment on the posterior region as in 

normal specimen. 


4. On the Azygos Veins in the Mammalia. By Frank 
BE. Brpparp, M.A. (Oxon.), F.R.S., Prosector to the 

[Received February 1, 1907. | 

(Text-figures 62-73.) 

(1) Introductory, p. 181. (9) The Azygos and other thoracic veins 
(2) The Azygos Veins in the Ungulata, | i NG vous ae UI gojpouciems 
p. 183 coypu, p- 213. 
eee ee 196 (10) The condition of the Azygos and 
(3) Insectivora, p. ; | the Classification of Mammals, 
(4) Lemurs and Apes, p. 197. p. 218. 
(5) Edentata, p. 198. | (11) The position of the Azygos with 
pel RO A | reference to the Vertebre, p. 219. 
a oes a ne (12) The Ven Intercostales Supreme, 
ar als, p. 2UU. 92 
2 p- 221. 
(8) Rodents, p. 208. (13) Conclusions, p. 222. 

(1) Introductory. 

It appeared to Cuvier*—and presumably to his editor Duvernoy, 
since the statement is left unannotated—that the Azygos veins in 
mammals were too variable to offer zoological characters of value ; 

* Anat. Comp. ed. 2, t. vi. (Paris, 1839) p. 238, 

182 MR, F. E. BEDDARD ON THE | Feb. 19, 

for he wrote, “ L’insertion de azygos, Vexistence d’une azygos du 
coté gauche sont assez variables; mais on sait que les mémes 
circonstances varient dans VPhomme. Elles ne méritent pas con- 
séquemment de nous arréter.” Muilne-Edwards*, on the other 
hand, and rightly, treats these veins as of importance, and tabu- 
lated the main variations. These statements really express the 
main facts as we know them today. With reference to the 
Marsupials, indeed, the assertion of Milne-Edwards seems to me 
to be nearer to the truth than some generalisations made more 
recently. This group is placed by Milne-Edwards under the 
heading “ Les deux veines azygos également développées.” 
Although I shall show reasons for slightly criticising this state- 
ment, it is not greatly exaggerated. On the other hand, 
denying the existence of a right azygos (“ Point de veine azygos 
«1 droite”) in the Sheep, Ox, Goat, Chevrotain, Pig, and Tapir he is, 
in my opinion, not by any means so accurate. With regard to 
the Tapir there is some error in Milne-Edwards’s statement ; for 
he also places that animal under the heading of those animals 
which only possess a right azygos. Sir Richard Owen’s classical 
text-book, published ten years later than the volume of Milne- 
Edwards's great work, adds but little to the record of facts con- 
cerning the azygos veins. There are, however, numerous scattered 
references to the condition of these veins in various mammals by 
Owen and others, to some of which I am able to refer in the 
course of the following pages. 

Many of these papers are quoted by Hochstetter? in his 
memoir dealing with the development of the Azygos (and other 
veins) in the Mammalia. A large number, however, relate to 
the condition of the azygos in man, and I do not attempt here to 
follow up that very large subject. I limit myself to such other 
mammals as I have been able to dissect, in many of which the 
azygos has not been described. The classificatory importance of 
the azygos has been recognised by Dr. Max Weber {, and there is 
no doubt that its conditions are often distinctive of genera or of 
whole orders of mammals. I propose, however, to deal with this 
matter after exposing the facts which [ have gathered together 
by degrees during several years of intermittent work upon the 
subject, which is a larger collection of facts concerning this vein 
than has previously been br ought together. 

Opinion with regard to the morphological nature of the azygos 
veins has lately undergone some change. Until lately the pre- 
valent view was that one or both of the postcardinals persisted as 
the Azygos, the Hemiazygos, or both. This view is embodied in 
diagrams in many text-books. 

Of recent writers Messrs. Parker and Tozier§ appear to hold 

* Anat. et Phys. Comp. vol. iii. (Paris, 1858) pp. 595-598. 

+ Morph. Jahrb. xx. 1893, p. 642 &c. Milne-Edwards’s résumé is largely based 
upon the observations of Bardeleben (Arch. f. Anat. u. Phys. 1848) and Marshall 

(“ Development of great Anterior Veins &c.,” Phil. Trans. 1850). 
t Die Saugetiere, Jena, 1904. 

§ “Postcardinal Veins in Swine,” Bull. Mus. Comp. Zool. vol. xxxi. 1898, p. 133. 


much the same view. For they remark that in the Pig “the 
hemiazygos” (which is the left azygos of my nomenclature) ‘‘ from 
the region of the heart to the tenth rib is therefore to be 
regar ded as the persistent anterior portion of the left post- 
cardinal.” The rest of the vein is formed, as they think, from 
“the accessory veins,” which appear to be the subcardinals of 
McClure*. They thus agree with Rathke in holding that the 
anterior part of the azygos is persistent posteardinal, but differ 
from him as to the mode of formation of the posterior region of 
the azygos; for Rathke held that this region was due to a con- 
tinual longitudinal anastomosis between intercostal veins, and 
was thus an entirely new structure. Zumstein tT put forward the 
older view also in the case of man, where that anatomist believed 
that he had traced the azygos and the hemiazygos to the post- 
cardinals exclusively. In the Guinea-pig, however, he = found 
that the posteardinals took practically no share in the formation 
of the azygos. Hochstetter $ came to conclusions which were not 
dissimilar. He allowed in the case of the Rabbit and of the Cat 
that the azygos of the adult down to about the eighth thoracic 
segment was the posteardinal, but that thereafter it was a new 
structure not formed from the postcardinal vems: “von da an 
caudalwirts aber ist sie eine Neubildung.” This region of the 
posteardinal, in fact, becomes a part of the posteaval. a pertectly 
different origin of the azygos veins is asserted by McClure || of 
Didelphys. ‘Excepting just at their entry into the duct of Cuvier 
they are quite independent of the postcardinals and of the sub- 
cardinals, though connected with the former by cross anastomoses. 
T shall bring forward various facts in the following pages which 
bear upon this question of the morphological nature of the azygos 
veins in mammals. 

(2) The Azygos Veins in the UNGULATA. 

I have paid special attention to this group since I have par- 
ticularly favourable opportunities, as compared with those enjoyed 
by other zoologists, of examining recently dead specimens. ‘These 
bulky animals obviously cannot be preserved, and must therefore 
be studied immediately after death. It thus follows that I am 
able to add a good deal to what is known upon the subject. 
It will be seen, however, that there is a very general agreement 
among the Artiodactyle division of that Order as contrasting 
with ‘the Perissodactyles, but that the division, as shown by 
the azygos vein, is not absolute. Max Weber, in lis recent text- 
book (Die Siugetiere, p. 642), uses the condition of the azygos 
to define the Artiodactyla thus :—‘ Die Vena azygos fehlt; die 

* “ Development of Veins of Didelphys,” Amer. Jour. Anat. v. 1906, p. 163. 
ii “ Bntwicklunge des Venensystems des Menschen,” Anat.-Hefte, Bd. vi. 1896. 
+ “ Venensystem bei dem Meerschweinchen,” ibid. Bd. vin. 1897. 

§ Loe. cit. p. 574. || Loe. cit. p. 185. 

184 MR. F. E. BEDDARD ON THE [Feb. 19, 

Vena hemiazygos miindet direkt oder indirekt in die Vorkammer.” 
But, as I shall show, this is not quite universal. 

Since the system of azygos veins is as perfect in the Gnu, 
Connochetes g grit, as in any other Ungulata, and more elaborately 
developed than in many, I shall commence with a description of 
those veins in this Antelope. The anterior vena cava receives a 
right and a left azygos vein which enter it very nearly, if not 
exactly, opposite to each other. Of these two the right is rather 
longer than the left. This right azygos receives five branches, of 
which the most anterior is composed of affluents from two ribs. 
It is evident therefore that the right azygos does not reach back 
nearly as far as the diaphragm. The left vein is composed of 
only four intercostal branches. The blood from the ribs lying 
behind these four ate connected, however, into another lon- 
gitudinal trunk lying on the left side. Seven or eight of these 
branches coming from as many intercostal spaces combine to 
form a vein running in the same straight line as the left azygos, 
but not joining it anteriorly. The vein, in fact, opens inde- 
pendently into the right auricle, as previous observers have noted 
for other Ungulates. 

It is not a little remarkable that the other species of Gnu, 
viz. Connochetes taurinus, which I have also dissected, shows 
differences in respect of these veins from Connochetes gnu. On 
the right side the azygos is a trifle smaller and collects blood only 
from four intercostal spaces. On the left side the azygos is very 
much longer, with seven or eight affluents. It enters the vena 
cava superior as in Marsupials &e., which possess two azygos veins. 
I did not detect any branch putting this vein into direct com- 
munication with the right auricle, such as occurs in Connochetes 
gnu and in some other forms which I shall deal with immediately. 
An intermediate condition is offered by another Antelope, Bubalis 
caama. In this animal the right azygos is composed of five 
affluents. On the left side we have both the anterior and the 
posterior vein of Connochetes gnu, and, as in that animal, the 
latter communication with the auricle direct. But a slender twig 
connects the two veins, which are thus a continuous vessel as in 
Connochetes taurinus. 

The azygos veins of Rhaphicerus melanotis present the usual 
Antilopine characters; but there are differences of detail from 
those of some other forms. On the right side, the internal 
mammary vein is followed in passing towards the heart by a 
superior intercostal which divides into two branches, each of which 
supplies the first or the second intercostal space. Behind this 
arises the right azygos, which is composed of three afiluents ; the 
first lies in the intercostal space 2/3, the others in 3/4 and 4/5. 
Thus the intercostal space 2/3 has two veins. On the left side 
there is a superior intercostal opening exactly opposite to the right- 
hand vein; but it isa larger vessel though it only draws blood from 
two intercostal spaces. The left azygos, as 1m alhed forms, 1s well 
developed and enters the heart in common with the vena cava 


inferior. It does not, however, consist only of a region lying 
behind its point of entrance into the auricle ; but a branch also 
runs forward supplying four intercostal spaces, the first being that 
between ribs 2/3. The vein does not join the superior intercostal. 
But it will be noted that rib interspace 2/3 has on the right two 

Ourebia nigricaudata is much like Rhaphicerus. The vena cava. 
anterior receives a pair of veins, the superior intercostals, which 
bring back blood from the region of the first rib only. The two 
veins are not absolutely symmetrical as to their point of entrance 
into the vena cava. The second to the fifth rib inclusive supply 
four intercostals which unite to form the right azygos. On the 
left side of the body the azygos consists of an anterior and a 
posterior section which unite at their communication with the 
right auricle. The anterior vein is very slender, excepting in the 
region at and near to its fusion with the posterior vein. The 
azygos proper, 7.e. the posterior section, is a stout well-developed 
vein which receives blood from the intercostals on both sides of 
the body, It ends at the diaphragm in a bifurcation formed by 
the two intercostals supplying the last (the thirteenth) ribs. 

Cephalophus grimmii showed an identity of arrangement in two 
examples, one a male, the other a female. The better developed 
left azygos entered the heart in company with the vena cava 
inferior, and not by a separate orifice. On the right side of the 
body the azygos consisted certainly of four intercostal affluents. 
The last two of these (at any rate in the male example) encircled 
the fourth rib. C. maawelli shows much the same relation to 
C. grimmii that the two species of Gnushow. For the left azygos 
opens into the precaval. But it gives off a branch to the auricle. 

Oryx leucoryx has also a comparatively short right azygos 
composed, however, certainly of five intercostal branches. The 
longer left azygos enters the heart either directly or with the 
vena cava inferior. Above the influx of the right azygos is a 
single inferior intercostal on the right side of the body, and on 
the left side a corresponding vein which, however, immediately 
divides into two trunks. There was, however, no connection 
between the posterior of these two veins and the lower section 
of the azygos system. 

The arrangement of these veins is much the same in Oryx beatria 
as it isin Oryx leucoryx. The left and principal azygos commences 
with the interspace between ribs 6 and 5. Above this is a vena 
suprema intercostalis, composed of only two branches lying in the 
first two rib interspaces. To this corresponds exactly on the right 
side a vein which draws blood only from the first intercostal space. 
Behind this the right azygos opens into the precaval opposite to 
the second or third rib. Its branches extend back to the sixth 
rib, and they begin with the second intercostal space. 

I have selected for figuring here (text-fig. 62) the azygos and 
immediately related veins of Cervicapra bohor, on account of their 
splendid condition in the male example which U had the opportunity 

186 MR. F. E. BEDDARD ON THE | Feb. 19, 

of dissecting in the month of December of last year. I am pretty 
certain that I have been able to note down the characters of all the 
veins of this system in the present species. The animal, I may 
remark, was young and very slightly diseased. Cervicapra bohor 
agrees in essentials with other Antelopes; but there are differences 

Text-fig. 62. 



Scl.m Vg ace 

IL /2 A 







Azygos veins of Cervicapra bohor. 

Az.l. Left azygos; Az.r. Right azygos; 7.m. Internal mammary; Sc/Z. Subclavian ; 
S.i. Superior intercostal ; p.c. Precaval. First three ribs numbered 1, 2, 3. 

of detail which are worth recording. The main azygos trunk lies 
as usual on the left side and opens into the heart. The level at 
which it bends towards the heart is opposite to the seventh rib. 


Its affluents, however, commence on the posterior side of the fourth 
rib. Beyond the last pair of affluents, which lie behind the 
thirteenth (and last) rib, the trunk is continued back as a very 
slender vessel. The first intercostal artery which crosses this 
azygos lies behind the tenth rib. Anteriorly, the second and 
third intercostal spaces give rise to vessels which unite into a 
trunk which debouches into the left subclavian vein. The right 

Text-fig. 63. 

Sel. pe. r7r7e- 
= dj i) See. 
eas ae 
= He : oe 
Aer e 
; | ueere Az.t 


Azygos veins ot Gazella euchore. Lettering as in text-fig. 62. 

subclavian, it may be remarked, joins the precava quite a con- 
siderable distance behind the opening of the left subclavian—a 
marked asymmetry which is not common. It receives, however, 
no branches from intercostal spaces and enters the thoracic cavity 
in the usual position—that is, in front of the first rib. Theright 
azygos is of fair size; it enters the precava about opposite the 
fourth rib. It receives branches from intercostal spaces 2 to 6 

188 MR. F. E, BEDDARD ON THE [Feb. 19, 

In Gazella euchore (the example dissected was a male) the dis- 
position of the several veins of the azygos system was as follows 
and as is shown in the accompanying illustration (text-fig. 63). 
The illustration in question shows that the subclavian veins are 
symmetrical with each other and occupy the usual position, 
emerging as they do in front of the first rib. To the left sub- 
clavian is attached the corresponding vena intercostals supreméa. 
This vein collects blocd from the second and third intercostal 
spaces. I am not absolutely certain whether its longitudinal 
trunk is not continuous with the left azygos. The latter com- 
mences with a branch from behind the fifth rib; it enters the 
heart on a level with the sixth or seventh rib. The right 
azygos 1s fairly well developed and enters the precaval vein on a 
level with the second rib. It collects blood by branches from the 
second to the fifth intercostal spaces inclusive. 

Tn a female Capra megaceros the left azygos also opened directly 
into the auricle; its first branch arose from behind the sixth rib. 
Anteriorly, a superior intercostal consisted of two branches 
lying respectively between ribs 1 and 2 and 2 and 3. On the 
opposite side a vessel corresponded exactly to this in point of 
entrance into the precava, but drew blood only from the first 
intercostal space. Immediately behind this opens the right LEA EOS, 
which is made up of four intercostal branches from ribs 2 to 5. 
In another example, also a female, the azygos was also on both 
sides, but the right vein was smaller than in the specimen just 
described and only drew blood from two intercostal spaces. 

Hemitragus jemlaica ( ¢ juv.) is rather different in the arrange- 
ment of these various veins. There is, on the left side, the typical 
Artiodactyl azygos entering the heart directly and commencing 
with a branch in intercosta i space 4/5. There is no superior inter- 
costal on this side; and on the right side there is only a single 
vein and that corresponds to the superior intercostal, since it 
draws blood only from behind the second rib. 

Of Nemorhedus swetienhami I have dissected a single male 
example. As usual, the left azygos was the predominant vein 
concerned with the intercostal circulation. It receives branches 
from the fifth rib onwards, and the main trunk was traced some 
way into the lumbar region behind the last (7. e. the thirteenth) 
rib. The vena cava anterior receives the right azygos and 
corresponding vein exactly opposite to it on the left side. The 
former receives blood from the first six ribs and ends entirely at 
the level of the sixth rib. The corresponding vein of the left side 
has naturally fewer branches and ends at the fourth rib. I did 
not ascertain whether there was any connection between the 
anterior and posterior series of intercostal veins on the left side of 
the body. 

In an old example of Phacocherus cethiopicus (text-fig. 64) the 
azygos veins were rather different from those of the two species 
just described. The general plan was the same: that is, there is 
a long azygos vein on the left side. This vein started with an 


affluent from the intercostal space between ribs 5 and 6. As far 
as the twelfth rib the vein lay outside the intercostal arteries, 
which up to this pomt passed between the azygos vein and the 
vertebral centra. The intercostal artery arising from the aorta 
between the twelfth and thirteenth ribs passed to the outside of the 
azygos. Asa rule among mammals this point, where the azygos 
changes its position relatively to the intercostal arteries, is further 

Text-fig. 64. 
mB P.c. 


Azygos veins of Phacocherus ethiopicus. Lettering as in text-fig. 62. 

forward. Anteriorly the precaval vein receives two quite sym- 
metrically disposed veins, one from each side of the body, which 
convey blood from the more anterior intercostal spaces. ‘These 
veins, moreover, are not only symmetrical with each other in so far 
as regards their point of junction with the precava: they draw 
blood from exactly the same intercostal spaces. Each consists of 

190 MR. F, E. BEDDARD ON THE [Feb. 19, 

four branches supplying the first four ribs. Their point of 
opening is about opposite the second rib. 

In a young example of the African Red River-Hog (Poéamo- 
cherus africanus) the azygos was also limited to the left side of 
the body, as in Porcula salvania. In Potamocherus, however, 
this vein enters the heart as in Ungulates; while in Porcula it 
seems to have the relations of the azygos in Marsupials &c., where 
a left azygos 1s present. The difference is not really of great 
importance as indicative of an anomaly in one or the other of these 
two genera of Suidee; for we have in other Ungulates—e. g., in 
Connochetes gnu—a left azygos which is divided into two parts 
whereof one opens into the jugular and the other into the heart. 
One or other arrangement is found in each of the Suine genera 
Porcula and Potamocherus. Potamocherus shows the arrange- 
ment which was found in Sus scrofa by Hunter * and quoted by 
Owen in his ‘Comparative Anatomy’? in dealing with the 
azygos veins of mammals. 

In the Pygmy Hog (Porcula salvania) the azygos vein is only 
developed upon the left side of the body ; there was absolutely no 
trace of this vein that I could discover upon the right side. It is 
large and thick, and its branches are important and obvious with 
the exception of the first. This more slender branch divides on 
issuing from the main trunk into three twigs, of which two run 
on each side of a rib. After this there are seven branches, which 
are large and were turgid with blood in the individual examined. 
These branches correspond of course regularly to the ribs; but 
they supply eight intercostal spaces, since the last of them, in 
which the azygos ends, bifurcates over its rib. The azygos Ties 
very definitely upon the left side of the aorta ; it is not median in 
position as is so often the case with this vem. The vem ended 
in front of the diaphragm ; nor could I detect any branch, however 
thin, which continued on the main trunk behind the diaphragm in 
the direction of lumbar veins or of the postcaval or renal veins. 
These observations refer to one example only, which was a female. 

The Musk Deer (dZoschus moschiferus) presents us with a system 
of azygos veins like those of other Ungulates. On the left side 
the well-developed azygos enters the aur ‘icle directly. On the right 
side the small azygos consists of one vein only. In front of this, 
and on both sides of the body, is a superior intercostal. 

Dorcatherium aquaticum did not show, as might perhaps have 
been expected, an arrangement of a specially primitive character. 
It is much like other Ungulates. There is a small azygos 
on the right side of the body formed by three aftluents only. 
The azygos of the left side only communicates with the right 
auricle. It is as extensive as is usual among Ungulates and con- 
sists of many intercostal affluents. The first of these branches is 
made up of two tributaries. There is no anteriorly running 

* ©Hssays and Observations on Natural History &e.,’ arranged by Richard Owen, 
vol. 11. 1861, p. 124. 
+ Vol. iii. 1868, p. 555. 


extension of this vein such as oceurs in, for example, Rhaphicerus. 
It is interesting to note that Dorcatheriwm does not agree with 
its nearest ally, Vragulus, as the following account of the latter 
genus will show. 

Of Tragulus meminna (text-fig. 65) I have been able to examine 
two adults—one of each sex—and a young one, nearly ready for 


Ga Sc. 
4 SLE = 7 

iV | f | 2 
bf ‘ 

ART pe 







| | 
Post. C. [oS | 

Azygos veins of Tragulus meminna. 

post.c. Remains of posterior cardinal of right side (?). 
Other letters as in text-fig. 62. 

birth, taken out of the uterus of the female. In all three the 
azygos was present only on the right side and entered the precaval 
very near to the heart. The nearness to the heart seemed to me 

192 MR. F. E, BEDDARD ON THE [ Feb. 19, 

to be more marked in the two adults than in the feetus. In the 
male adult the first affluent of the azygos arose behind the fourth 
rib; after this there was a gap of one rib, and three very much 
stouter affluents arose behind the sixth, seventh, and eighth ribs 
respectively. Thereafter the psoas muscles concealed the inter- 
costal veins at their entry into the azygos, but they appeared to 
be regular. In the female Deer the same very stout intercostal 
veins arose from the intercostal spaces 6-7, 7-8, 8-9 as in the male, 
but I did not find the slender anterior intercostal which I have 
described in the male; I should not like, however, to assert that 
it was absent. It is very interesting to observe this constancy in 
a vein which has been stigmatised as most inconstant. Moreover, 
the foetus showed the same arrangement of these veins with a 
slight difference. The first affluent of the azygos was very stout 
and arose between ribs 3 and 4, being thus a,rib further forward ; 
but after this the first affluent of the regular series arose in the 
same way as in the two specimens of 7ragulus meminna Just 
described, behind the sixth rib. In this feetus, moreover, a very 
slender intercostal, which I did not find in either of the adults, 
arose from behind the second rib and either joined the precaval 
independently or just at the point of entrance of the azygos. In 
this fcetus the first rib sent an intercostal vein to the subclavian 
of the right side. On the left side none of the three specimens 
had any properly developed azygos. But in the foetus and the 
male adult a large cesophageal vein received intercostal affluents 
from the second and third ribs on the left side and entered the 
subclavian of that side. There was something to correspond in 
the female; but Iam unable to give a proper description. In any 
case, the presence of rather more veins 1m the fcetus and the absence 
of at any rate much variation in the azygos system of the adult 
Tragulus meminna *, ave noteworthy. It is possible indeed that 
there was no variation at all. 

There is a very considerable agreement between the azygos vein 
of Tragulus and that of Cervus sika, but not, as will be shown pre- 
sently, of Cervus aristotelis. Inafemale of Cervus sika the azygos 
was developed only upon the right side of the body. There were no 
traces of anything of the kind on the left side. The right azygos 
enters the precaval rather forward—in fact, opposite to the second 
orthird rib. The first affluent which joins the azygos runs parallel 

* An interesting note upon certain points in the venous system of this Deerlet 
(“The Posteava of an adult Indian Chevrotain, Tragulus meminna Krxieben,” 
Anat. Anz. Bd. xxix. 1906, p. 375), by Prof. McClure, has come into my hands 
through the kindness of the author during the writing of the present memoir. In 
this note it is remarked that, contrary to what is found in most mammals, but 
agreeing with the conditions observable in Dasypus, Elephas, and the Marsupials 
(generally), the postrenal division of the postcava lies directly ventrally to the aorta, 
instead of dorsally and to one side. Dr. McClure naturally wonders if this is cha- 
racteristic of Tragulus meminna, or is abnormal. I looked into the matter carefully 
in the three specimens upon which I have reported above, and find that Dr. McClure 
has discovered a perfectly normal character of this primitive Ruminant. I may 
furthermore point out that he figures (Joc. cit. fig. 2) the rigbt renal vein as double 

and the left as being single as well as much longer. I found this also to be the case 
in the adult raale and in the fcetal male, but not in the adult female. 


with it for some little distance as a longitudinal trunk, and receives 
branches from the second to the fifth mtercostal spaces inclusive. 
The main trunk of the azygos commences with an affluent from 
the sixth intercostal space. There is, it will be observed, an 
obvious likeness here to certain Rodents also; for instance, 
Celogenys*. I may also observe that the vein lettered post.c. in 
text-fig. 65 also probably corresponds to this additional azygos, for 
the nature of which see p. 213 below. 

In Cervus aristotelis the azygos on the right side of the body 
pours blood into the single precaval from five aftluents, which 
collect. blood from the first five intercostal spaces. On the left 
side there is a corresponding vein which, however, flows into the 
precaval rather in front of the right azygos, between the second 
and third ribs. This collects blood from the first and a few sub- 
sequent intercostal spaces. I think that it is continuous with the 
usual Artiodactyle left azygos, which enters the heart opposite to 
the sixth rib and extends back to the diaphragm. 

Of Perissodactyle Ungulates I have only examined Lquus 
chapmani. The arrangement here was precisely as in the Horse. 
The azygos was present upon the right side only. The first 
affluent contributing to the vein arose from the interspace 
between the fifth and sixth ribs, and the vessel appeared to end 
posteriorly after the affluent from the thirteenth rib. 

Hyrax capensis.—This animal shows affinities to the Ungulates 
in the structure of the azygos veins combined with a good many 
differences from such Ungulates as I have been able to examine. 
It is noteworthy that this ‘‘Subungulate” agrees with the Horse 
and Tragulus vather than with most Artiodactyles in the much 
greater size of the right azygos. In view of other comparisons 
that have been made between Hyrax and the Perissodactyle 
Ungulates, this additional fact is of interest. In the Rhinoceros, 
according to Owen *, the right azygos is the principal azygos. This 
feature alone is, however, “obviously not enough to establish an 
affinity with the Per issodactyle section of the Ungulata, for many 
mammals possess only the right azygos, with or w ithout traces of 
the left. It is the characters of the azygos on the left side which 
indicate the Ungulate affinities of this animal. In one of two 
examples of Hyraa capensis which I have dissected, the first two 
costal interspaces were occupied by veins which united to form a 
single trunk opening into the vena cava anterior. These represent 
the superior intercostal vein of other mammals. On the right side 
of the body the corresponding veins were present, but each opened 
separately into the vena cava. The next two ribs on the left side, 
2. e. Nos. 3and 4, were supplied by a vein each, the two veins lying 
both of them in the same intercostal space, between ribs 3 and 4. 
These united to form a slender trunk which passed backwards 

* See p. 209. 
+ “On the Anatomy of the Indian Rhinoceros,” Trans. Z.S. iv. p.46. The Tapir 

also has been stated to possess a large right azygos. 

Proc. Zoou. Soc.—1907, No. XIIT. 13 

194 MR. F. E. BEDDARD ON THE [ Feb. 19, 

and opened into the vena cava inferior (text-fig. 66) just before 
the debouchment of the latter into the right auricle. This is 
clearly the Artiodactyle condition of the vein in question, though 
differing in detail from that of any particular Artiodactyle that 
has been studied and described. For in that group there is either 

Text-fig. 66. 

aeons) (EE ae ees: 

LT Tre 


Hyrax capensis; two different arrangements of azygos system. 

post.cav. Postcaval. Other letters as in text-fig. 62. 

an anterior and a posterior branch of the left azygos which unite 
close to the opening into the auricle or vena cava posterior, or the 
posterior branch alone is present. In Hyrax it is the anterior 
branch alone which is present. 


The corresponding ribs on the right side were supplied with 
veins joining the long right azygos. The azygos system of Hyra« 
has been partly described by Brandt* in a memoir upon the 
general anatomy of Hyrax, who observes that the vena cava 
posterior enters the right auricle ‘ nachdem sie die vorderen 
Vene intercostales und die Vena azygos aufgenommen.” ‘The same 
fact, according to Weber 7, has been also noted by George =; but 
I have not had the opportunity of studying this paper §. Nothing 
is said by Brandt concerning the right azygos. It is, I presume, 
legitimate to compare the descending region of the left azygos in 
Hyrax with the left anterior cava. In a second specimen I did 
not find a left azygos opening into the vena cava inferior. But 
JT am not inclined to deny its existence. The large right azygos 
was equally well developed in this example. 

In a third example of Hyrax capensis (a female) the azygos 
system was entirely developed upon the right side of the body, 
and differed considerably from that of the two individuals just 
described. The main azygos stem (text-fig. 66) was traceable 
to the lumbar region, where it communicated with the vena cava 
inferior (not opposite to the renal vein) in common with a lumbar 
vein. The vein undoubtedly belongs to the right side since it 
lies on the trachea on the right side of that tube. Its mode of 
termination anteriorly is, so far as my own experience goes, unusual 
among mammals. Instead of entering the jugular some little 
distance in front of the heart, as is elsewhere (so far as | have seen 
in my own dissections) invariably the case with the right azygos 
when present, it enters that vein so near to its entry into the 
auricle that it may almost be said to enter the a 
But there is, of course, no question as to a direct communication 
with the coronary sinus, like the left azygos of the Cavicornia. In 
front of this. vein, which debouches into the heart opposite to the 
fifth rib, is a small vein which receives blood from the 2nd and 
3rd intercostal spaces. 

The disposition of the azygos veins in the Artiodactyle Ungu- 
lates seems, therefore, to be fairly constant in the group, though 
the left azygos does not quite invariably open directly into the 
right auricle or into the vena cava posterior just before the open- 
ing of the latter into the right auricle. This arrangement of the 
left azygos has not been found in any other group of mammals 
excepting only in the Mole, where it has been stated to be the 
same as in the Artiodactyles. Possibly this fact may be con- 
sidered as requiring confirmation. It is, however, interesting to 
note that as an abnormality the left azygos in man may open 
into the right auricle directly. I have not attempted to compare 

* Mém. Ac. St. Pétersbourg, ser. 7, t. xiv. p. 65. 

+ Die Saugetiere, Jena, 1904. 

+ Bibl. Ecole Hautes-Htudes, Sec. Sc. Nat. xii. 1875. 

§ In an earlier paper by George (Ann. Sci. Nat. (6) 1. 1874) a right azygos only is 

described. : J 
|| This is also the case with the Horse, according to Chauveau and Arloing. Ihave 

also described the same arrangement in Zragulus. 

196 MR. F. E, BEDDARD ON THE | Feb. 19, 

the variations in the human subject with the conditions which 
are normal in other mammals, and it is possible that many 
examples have been recorded of this particular abnormality in 
man. I refer only to one instance, which I cull from the list of 
literature given by Hochstetter in his memoir upon the develop- 
ment of the veins in mammals*. 


My notes upon this group are very few, but not without 
interest. Milne-Edwards has observed + that the Mole possesses 
both azygos veins, and that the left debouches into the right 

Text-fig. 67. 



a a ae oes 
= {=e fe 
We ee 






Azygos veins of Hrinaceus algirus. Lettering as in text-fig. 62. 

auricle precisely as in certain Ungulates. This fact is not referred 
to by Owen, who however states + that “the left vena azygos 
communicates with the left precaval in the Hedgehog and many 

* Gruber, “ Ueber einen Fall von Einmiindung der V. hemiazygos in das Atrium 
dextrum cordis beim Menschen,” Arch. f. Anat. u. Phys. 1864, p. 729. 

+ Anat. et Phys. Comp. 11. 1868, p. 595. 

+ Comp. Anat. & Phys, Vertebrates, vol. ii. 1868, p. 553. 


others [of the “ Lissencephala”], and is larger than the right.” 
This does not exactly agree with what I have found in an example 
of Hrinaceus algirus, 9. Two azygos veins were present (text- 
fig. 67) and of equal length. Each opened into the precaval of its 
own side at a point corresponding to the interval between the 
second and third ribs. In the ease of the right azygos, the first 
affluent arising between the second and third ribs and the second 
arising between ribs 5 and 6 were particularly large. It is note- 
worthy that posteriorly there was also an irregularity in the 
branches of the azygos, one branch often serving two intercostal 
spaces. The specially thick affluents were not noticeable on the 
left: azygos. 

I have also dissected one example of the Common Hedgehog 
(Erinaceus ewropeus), which differs both from Owen’s account of 
that species already referred to and from Hrinaceus algirus as 
dissected by myself. In this Hedgehog the right azygos alone was 
well developed, and commenced with an affluent from the first rib 
space, opposite to which it entered the right precaval. On the 
left side a very small azygos was present consisting, so far as I 
could see, of only one branch arising from the first intercostal 
space. It joined the left precaval at a point apparently exactly 
opposite the right azygos. 

(4) Lemurs and APES. 

Of this group I have examined a considerable number of species, 
and in all of them the right azygos as a distinct vein is present, 
and alone present. I have found this to be the case in the 
following, viz. Lemur catta (3 examples), L. mongoz, L. xantho- 
mystax, L. macaco, L. albifrons, L. coronatus, L. varius, Nycticebus 
tardigradus (3 examples), Perodicticus potto, Galago crassi- 

Although it is correct to say that there is only a right azygos 
in these Lemurs as a complete vein entering the vena cava anterior 
on its own side of the body, there are in a few forms traces of the 
left azygos—of a hemiazygos. Thus in Galago crassicaudate there 
is a longitudinal trunk springing from the right azygos opposite 
to the 7th intercostal of the right side of the body. This vessel 
runs forward, receiving intercostals from the left side and finally 
enters the left subclavian. This vein is obviously the left superior 
intercostal of man, but is more extensive in Galago than in Homo. 
Furthermore, a posterior fragment of the same vein and not con- 
tinuous with it is left in the shape of a longitudinal connection 
between the 8th and 9th intercostals of the same side of the body. 

In Lemur albifrons I found a hemiazygos arising from the right 
azygos shortly after the origin of the first of its intercostal 
branches, which crossed to the left side and received the inter- 
costal veins of that side of the body. 

Of the higher Primates I have examined a considerable number 
of species, in all of which there is but one azygos, the right, with, 

198 MR. F, E. BEDDARD ON THE [Feb. 19, 

in cases, a left hemiazygos. I do not for the present give any 
details concerning the Monkeys and Anthropoid Apes. 

(5) EpENTATA. 

Of this group of mammals I have only been able to examine a 
few of the South-American forms. 

In the Great Anteater, I/yrmecophaga jubata, there is only a 
right azygos, with no traces that I could discover of the left-hand 
vessel. As this statement refers to three examples, including 
both sexes, it 1s probably a statement of the normal state of affairs 
in this animal. In one example, at any rate (I have not notes on 
the others), the first affluent occupied the 4th intercostal space. 

Of the small Anteater, Zamandua tetradactyla, | have examined 
but a single specimen, in which the conditions of the azygos were 
quite the same. 

In two Armadillos, Dasypus villosus and D. minutus, the right 
azygos was also alone present. Hyrtl, in his account of the 
anatomy of Chlamydophorus, only found, or at least only men- 
tions, the right azygos. These facts afford an additional argu- 
ment, though doubtless a small one, for the banding together of 
the American Hdentates. 


The azygos vein in this order of mammals presents a very 
uniform arrangement. I have dissected a considerable number 
of species belonging to many genera, and in the great majority of 
them there is but one azygos present, which is that belonging to 
the right side of the body. In these I have not been able to find 
any trace of the vessel of the left side. The following species 
present this condition of the azygos veins, viz. :—Galictis barbara, 
Crossarchus obscurus, Cynictis levaillanti, Viverra civetta (2 ex- 
amples), Vandinia binotata, Cercoleptes caudivolvulus (3 examples), 
Nasua rufa (2 examples), Felis pardus, Procyon cancrivorus 
(3 examples), P. lotor, Lutra vulgaris (2 examples), Helictis per- 
sonata, Cryptoprocta ferox, Herpestes griseus, H. pulverulentus, 
Arctogalidia trivirgata, Proteles cristatus, Gulo luscus. 

In two specimens of Procyon cancrivorus the azygos of the 
right side received a hemiazygos from the left side about halfway 
down, and there were no vene intercostales supreme. In Procyon 
lotor I found no hemiazygos, but there were two vene intercostales 
superior on the right side, supplying the first two ribs and opening. 
separately into the precava. On the left side a single vein opened 
into the subclavian. 

The azygos vein in the Hyena (Hyena crocuta) (text-fig. 68) is 
the most remarkable modification of this vein which I observed 
in the Carnivora. As has been noted, the azygos in the Carnivora 
is constantly a long S-shaped vessel lying on the right side and 
extending down to the diaphragm, giving off regular branches. In 


this Hyena, however, the azygos is singularly short, supplying 
only three intercostal spaces. It opens into the Ductus Cuvieri by 
a single stout branch ; this is formed by the union of three branches 
which are of very unequal calibre. The most anterior of these is 
a moderately stout vein running slightly forward in direction. 
The middle vein of the three is very markedly the stouter. It 
plunges at once, after the shortest possible course, into the thick- 
ness of the parietes. The third branch is more slender than both 
the first and the second. It also has a very short course obliquely 
backwards, and is very soon lost in the parietes. it gives off one 
branch to the left side and a slender short main azygos. I could 
discover no vein upon the left side of the body. It is to be 
remarked that we have here an abortive azygos, the first branches, 
or perhaps branch, only being well represented. These collectively 
correspond, as I think, to the first branch of the azygos in some 
other forms, which is very frequently subdivided into two or even 
three branches. 

Text-fig. 68. 

Azygos of Hyena crocuta. Lettering as in text-fig. 62. 

It is usual for the first branch of the azygos to arise between 
the fifth and sixth ribs in this order of mammals. I have found 
this to be so in Procyon lotor, Lutra vulgaris (one example; in 
another it arose between ribs 4 and 5), Swricata tetradactyla, 
Genetta felina. 

In Crossarchus obscurus (text-fig. 69) the azygos entered the 
precaval about opposite to the fifth rib. Its most anterior affluent 
arose between the third and fourth ribs. Between the first and 
second ribs an intercostal arose on the same side (the right) and 

200 MR. F. E, BEDDARD ON THE [ Feb. 19, 

Text-fig. 69. 

Scl-= - Sel. 
Qy | 7 
St mb sa pails 

al ( 

“a | 

AZ rE 

Azygos of Crossarchus obscurus. Lettering as in text-fig. 62. 

entered the subclavian vein. The corresponding vein of the left 
side of the body was formed of two aftluents arising from the first 
two intercostal spaces, which opened into the precaval and noé into 
the subclavian. 

(7) MarsupPIALs. 

T am able to give rather a fuller account of the genera of this 
group than of the other groups of mammals treated of in the 
present communication. And furthermore, of several genera 
which I have examined I have had the opportunity of seeing 
several species, and occasionally more than one example of the 
same species. The azygos veins of Marsupials have been lately 
the object of a careful study by Prof. McClure * in connection 
with a detailed survey of the blood system of the Opossum, 
Didelphys marsupialis. He has in his memoir collected together 
what has been written upon the subject of this vein, and I 
shall review his conclusions in the light of the additional facts 
which Iam here able to contribute to this branch of anatomy. 

* American Journ. Anat. 11. 1903, p. 371. 



T shall commence with the Kangaroos, of which I have examined 
a good many different species. 

In Macropus wualabatus the condition of the azygos veins is the 
most primitive of that to be seen in Marsupials. There are two 
veins, one on each side, each, of course, opening into its corre- 
sponding vena cava anterior. These veins appear to be exactly 
equal in length. 

In Macropus agilis I have found one of the two opposite 
extremes. In the single example of this species which I have 
dissected there was but one azygos, lying upon the right side of the 
vertebral column. In the preceding species of Macropus the two 
azygos veins end, or at any rate dwindle to almost nothing, in the 
neighbourhood of the last rib behind the diaphragm. Their last 
affluent is derived from the body-wall close to the last rib. In 
the specimen of JMacropus agilis, on the other hand, the single 
right azygos is continued as a wide vein for a long distance back- 
wards. <A little in front of the kidney it ceases to be superficial 
and is imbedded in the dorsal musculature. It then emerges and 
gives off a branch joining the vena cava posterior in the region 
of the kidney. This vessel is not the renal vein itself, for the 
latter can be followed from the kidney to the vena cava as a 
distinct and much thinner vessel. Behind this again the azygos 
dies away towards the pelvis, retaining till its extremity a con- 
siderable size. 

In Macropus melanops I found in one individual the same state 
of affairs as in the last species in so far as only a single azygos 
was present, and that on the right side. This vein, however, was 
slender and drew blood from only six intercostal spaces, after 
which it dwindled or entirely disappeared. In a second specimen 
of the same species there was also a limited azygos on the right 
side ; but, in addition to this, traces of one belonging to the left 
side of the body, which, however, was quite small and only drew 
blood from one intercostal space. 

An example of Wacropus bruni showed conditions intermediate 
between the two extremes already dealt with. In this species 
there was an azygos on the right side extending as far back as to 
the diaphragm. On the left side there was a less completely 
developed vein only drawing blood from four intercostal spaces. 

A female Macropus dorsalis was not very different. In this 
Kangaroo the azygos was only properly developed upon the right 
side of the body. It was prolonged, however, for a long way back 
as in Macropus agilis. In the same way a branch was given off 
to the vena caya posterior underlying the renal vein. The AZY Os 
trunk ended some way behind this. 

On the left side of the body there were traces of a left azygos. 
This consisted of the azygos proper, being a short vessel gathering 
blood from one intercostal space only, and of another trunk in 
front of this flowing separately into the vena cava anterior of that 
side of the body, which corresponds to the superior intercostal of 
other mammals. 

202 MR. F. BE. BEDDARD ON THE [Feb. 19, 

A male Macropus giganteus showed an almost exactly similar 
arrangement of the several azygos veins. On the right side the 
vein was strongly developed and passed back to the pelvic region, 
giving off a branch to the vena cava posterior in the region of the 
kidney. On the left side one vein arose from the vena cava 
anterior sinistra which apparently corresponds to the two which 
arose separately in Macropus dorsalis. This vein was formed of 
two afiluents only. 

A female Macropus rufus possessed a well-developed azygos 
upon the right side, which however did not extend backwards to 
anywhere near the pelvic region. I traced it to the eleventh rib, 
between which and the tenth lay its terminal branch. The vein 
supplied all the intercostal spaces between this and the fifth 
anteriorly. On the left side of the body wasa rudimentary azygos 
of two branches lying between the fourth, fifth, and sixth ribs. 
A second example, a male, showed an identical arrangement of 
both azygos veins. 

Macropus alligatoris is precisely like Macropus rufus. In a 
male specimen of this Kangaroo the well developed azygos lay 
upon the right side of the body, while on the left side was a rudi- 
mentary vein formed of two afiluents only. 

Exactly the same description is to be given of J/acropus 

Of Macropus derbianus I have had the opportunity of seeing 
three individuals. Two of these were of particular interest since 
they were mother and daughter. The female fcetus showed two 
azygos veins very nearly equal in extent, but the right was rather 
longer than the left ; this diserepancy—it will be observed—being 
the usual one. On the other hand the parent showed reverse 
conditions; the left azygos was longer than the right, and in show- 
ing this character was unique among the examples of this genus 
which I have examined up to the present. But ina third specimen 
of the same species, adult and a male, the azygos veins were as 
nearly as possible equal, the left being again a trifle the longer. 

In Halmaturus bennettii 1 found the azygos on the left side 
to be distinctly longer than that of the right side. JI counted in 
fact seven affluents on the left side. On the right side there 
were only five branches, of which the first followed thie fifth rib. In 
a second specimen (text-fig. 70), a male just out cf the pouch, the 
small left azygos had only three afiluents from ribs 4 to 6. The 
right azygos began with an affluent from rib 2. 

“Of the genus Dendr olagus | have examined only one species, viz. 
D. bennettii. In this Tree-Kangaroo I found the azygos to be 
present only on the left side. 

Apyprymnus rufescens is a species which shows differences in 
the proportions of the two azygos veins. In one example, a female 
the left azygos was shorter, composed of four affluents; the right 
reached to the diaphragm. Ina male the right was very much the 
shorter, composed as it was of only two affluents. On the left 
side there were four or five branches before the vessel disappeared 


into the thickness of the musculature. Later on this azygos 
dilated to form a thicker vessel which lay beneath the aorta and 
reached nearly to the iliacs. This is obviously quite comparable 
to what I have described above in Macropus giganteus and other 
species; but in them it was the right azygos which was thus 
prolonged and enlarged. J could not be certain whether in Zpy- 
prymnus rufescens there was or was not a connection with the 
renal vein or the postcaval. In this specimen there was also on 
the left side a superior intercostal vein flowing separately into the 
vena cava superior. 

Text-fig. 70. 


Azygos veins of Halmaturus bennettii. Lettering as in text-fig. 62. 

In a third example of Wpyprymnus rufescens, which was a 
female, the conditions observed differed from those of either of 
the other two. The two azygos veins were absolutely symmetrical 
so far as I could see, and moreover this symmetry extended to 
quite small details. On both sides of the body the last two of the 
four intercostal affluents of each azygos joined before pouring 
their contents into the azygos. It is curious to find here in the 
same species all the chief varieties shown by the azygos. 

While Zpyprymnus rufescens is an example of a Marsupial 
in which the condition of the azygos veins varies from individual 
to individual, the Common Phalanger (7richosurus vulpecula) 
offers precisely the reverse characteristic. Of this species I have 
dissected six examples belonging to both sexes. In all of them 
the azygos vein of the left side is fully developed and reaches back 
as far as the diaphragm. In all of them the right azygos is present 

204 MR. F. HE. BEDDARD ON THE [Feb. 19, 

but is invariably small, and consists at the most of a short trunk 
made up of affluents from three intercostal spaces only. There is 
some difference in the exact number of these aftluents; there are 
either two or three or apparently only one. In one example I 
recorded the further extension backwards of the left vena azygos 
till it reached the vena cava behind the kidney. This dis- 
position of the azygos agrees with what has been already recorded 
by previous observers concerning this species. These previous 
observations refer to several examples. The state of affairs may 
therefore with confidence be regarded as characteristic of the 
Allied to the Vulpine Phalanger is the Wombat (Phascolomys 
mitchell) (text-fig. 71), of which I have dissected two examples from 

Azygos veins of Phascolomys mitchelli. 

os. (Hsophageal vein. Other letters as in text-fig. 62. 

the present point of view. In the first example, a female, two 
azygos veins were present and about equalled each other in length. 
The conditions were in fact like those of Macropus ualabatus. 

In a second Wombat, also a female, the conditions of the 
azygos velus were apparently identical. Both veins were present 
and well developed; but the right-hand one was the longer 
of the two. In front of the two azygos veins no intercostal 


branches entered either of the two precavals. The right azygos 
entered the precaval at about on a level with the fourth rib. The 
first branch arose between the third and fourth ribs. The position 
of the opposite azygos was the same. The right azygos continued 
down to the tenth rib unaltered; at this point it divided into two 
branches, of which the outer supplied the intercostal spaces ; the 
inner branch, which was more slender, received at least one cross 
branch from the outer and could be traced some little way back as 
a very slender vessel; I did not ascertain its posterior connections 
ifany. Just after its bifurcation it was crossed by the first inter- 
costal artery to cross it, those in front lying below the azygos. 
This division of the azygos posteriorly reminds us of the figure of 
the Pig’s azygos given by Messrs. Parker and Tozier*. “But it 
seems more likely ‘that the inner branch is not the hemiazygos, 
but possibly the subcardinal of the right side, the outer branch 
being in that case the true persistent postcardinal. These matters, 
however, are more fully dealt with below. The left azygos of this 
example of Phascolomys mitchelli was shorter than the right; it 
ended absolutely between the tenth and eleventh ribs. 

In the Brush-tailed Rock-Wallaby (Petrogale penicillata) the 
disposition of the azygos veins was like that which is on the 
whole characteristic of the genus J/acropus. I have dissected 
three examples, one male and two females. They all agreed 
except in minutiz. In all of them the right-hand azygos was the 
one to be well developed. But I never found that this vessel had 
a prominent backward prolongation to the pelvic region as is 
occasionally to be seen in Macropus. On the left side there is 
only a trace of the azygos, and this is either derived from only 
one or from two intercostal twigs. In P. wanthopus, according to 
Parsons‘, the right azygos is also the larger, or rather the only 
one present. 

I do not know whether it is necessary to separate Bettongia 
peniciliata from B. ogilbyi. The condition of the azygos veins 
offers no help in deciding this question, since the two examples of 
B. penicillata differ considerably from each other, and I have only 
a single example whych came into my hands labelled Bettongia 
ogilbyi. In one specimen of Lettongia penicillata, a female, the 
right azygos alone was well developed, extending far back towards 
or to the diaphragm. On the left side the azygos was very short, 
consisting of two intercostal affuents only. In the second example 
of this species, which was also a female, the azygos on both sides of 
the body was about equally developed. In the single ettongia 
ogilbyi the two azygos veins were well developed; but that at the left 
side was distinctly the longer of the two. It is to be noted that if 
we are to unite these species, as Mr. Thomas has done in his 
‘Catalogue of Marsupials in the British Museum,’ the differences 
exhibited in respect of the azygos vein are almost exactly the same 
as those shown by “pyprymnus rufescens. 

* Bull. Mus. Comp. Zool. xxxi. p. 138, fig. 4 
+ See P. Z. S. 1896, p. 706. 

206 MR. F. E. BEDDARD ON THE [ Feb. 19, 

In a single example of Dromicia nana the azygos was well 
developed on the left side of the body and there was a short 
azygos on the right side. 

In a species of Pseudochirus the azygos was present on the left 
side only ; in addition to the azygos an anterior intercostal flowed 
separately into the vena cava anterior, deriving its blood from a 
single intercostal space only. In a second example (P. peregrinus) 
I found the same. The azygos proper commences with the second 
rib and joins the vena cava in the neighbourhood of the left kidney. 

A single male example of Petawrus breviceps, which was injected 
to illustrate the anatomical relations of these veins, possessed an 
azygos on the left side only. 

In the Thylacine the left azygos is the predominant one, and 
there is also a left superior intercostal vein entering the precaval 
independently of the azygos. The right azygos is, however, by 
no means rudimentary for it supplies four intercostal spaces. 

Phascologale penicillata has likewise a well-developed azygos on 
the left side extending back to the diaphragm. I saw none on 
the right side of the body. 

In a female Dasywrus viverrinus the azygos veins were almost 
exactly like those of the Thylacine. The left-hand vein was well 
developed, extending back almost as far as the diaphragm ; on the 
right the azygos was formed of five affluents. Inamale D. mauget 
the left azygos was well developed and there was no trace of a 
right-hand vein. 

A female Perameles obesula agreed with Phascologale rather 
than with the last species; for the azygos was present only on 
the left side of the body. 

Of Onychogale frenata I have examined a single female example. 
The azygos was predominantly developed on the left side of the 
body. It flowed into the jugular at a point about opposite to 
the fifth rib. The first branch of this left azygos formed the 
intercostal vein lying between ribs 4 and 5. Thence regular 
branches were given off as usual. The main stem of the vein 
became much thinner in the region of the diaphragm ; but there- 
after increased notably in volume and could be traced back down 
into the pelvic region as a massive vein of not much smaller 
calibre than the vena cava, alongside and to the left side of which 
it ran to a point which I did not determine. There is an obvious 
similarity in this case to that of Wpyprymnus rufescens described 
above. The azygos was connected to the vena cava by a stout 
branch in the neighbourhood of the kidney. An anterior vein, the 
superior intercostal, arose separately from the left jugular and 
supplied ribs 3 and 4, There flowed into the right jugular 
at a point corresponding to that of the left azygos a much 
smaller right azygos. This slender vein only drew blood from ribs 
4 to 7. 

The results obtained from the dissections enumerated in the 
foregoing pages enable me to revise some of the conclusions 
arrived at by Dr. McClure in his memoir referred to above. 


Dr. McClure is of opinion that “a single azygos vein is the rule 
in Marsupials, and that when two are present the case may be 
regarded as a variation.” The facts gathered by myself seem to 
me to show that the double azygos of the Monotremata is largely 
preserved in the Marsupials, so much so that the process of dis- 
appearance of one or the otheris but rarely completed. There are 
but few forms in which there is absolutely no vestige of either 
right or left azygos as the case may be. This is precisely what 
would be expected in view of the admittedly archaic position of 
the Marsupials, accepting, of course, the view now generally held 
that they are not intermediate between the Prototheria and the 
Placentals, but are an offshoot of an early Eutherian. For in 
the Rodents, an admittedly primitive type, we have considerable 
traces of the double azygos. This latter point, however, will be 
discussed in relation with the azygos in Mammalia generally on 
a subsequent page. On the other hand, my own observations 
confirm Dr. McClure when he remarks that a right azygos is 
characteristic of the genus J/acropus, and that a left azygos is 
characteristic of the Phalangeride. 

J would rather extend Dr. McClure’s remarks about the condition 
of the azygos vein in the Carnivorous Marsupials, and point out 
that in this whole group the existence of a prevalent left azygos 
is the rule so far as we know at present. The table given by 
Dr. McClure of the condition of the azygos in such Marsupials as 
were known when he wrote, brings out very clearly the variation 
in respect of the veins not merely from genus to genus but from 
species to species. This generalisation I confirm with great con- 
fidence, and also would emphasise the very frequent variation of 
these veins from individual to individual. The condition of the 
azygos in fact is by no means so fixed in this group as it is in the 
Carnivora, for example; and modification is evidently progressing 
along two lines, in ene of which the right azygos and in the other 
the left azygcs is being retained, while the opposite vein is in 
process of disappearance. 

Noteworthy, too, is the occasional large size of the postdia- 
phragmatic continuation of the azygos in the Kangaroos, Macropus 
giganteus, M. agilis, &e., which is, so far as my own observations 
go, invariably of the right azygos. 

The above results may be conveniently tabulated as follows :— 

Azygos on both sides equal or nearly equal. 

Macropus wualabatus M. bruni, MW. derbianus, M. bennettit, 
Phascolomys mitchelli, Bettongia ogilbyt, Eipypr ymnus rufescens, 
Thylacinus cynocephalus, Dasyurus viverrinus. 

6. Azygos large on right side, with a rudiment on left of not 
more than three intercostal affluents. 

Macropus dorsalis, M. giganteus, M. rufus, M. alligatoris, 

M. antilopinus, M. melanops, Petrogale penicillata, Bettongia peni- 

208 MR. F. E. BEDDARD ON THE [ Feb. 19, 

c. Azygos only present on right. No rudiment on left side. 
Macropus melanotis. 

d. Azygos large on left side, with a rudiment only on right 
Trichosurus vulpecula, Dromicia nana, Epyprymnus rufescens, 
Onychogale frenata. 

e. Azygos on left side only. No rudiment on right side. 

Dendrolagus bennettii, Didelphys nudicaudata, Pseudochirus, 
Perameles obesula, Petaurus breviceps, Phascologale penicillata, 
Dasyurus mauger. 

(8) RopeEnts. 

Of the azygos in this order of Mammalia Owen wrote that 
“opposite proportions [to those shown by the azygos veins of the 
Hedgehog] prevail in Leporidee and some other Rodents, as in 
Squirrels, the left azygos being small or wanting.” ‘This state- 
ment is largely but not entirely true, as will be seen in the course 
of the following description of several genera and species of 
Rodents. In view of the fact that so many Rodents possess, as 
is well known, both precaval veins, it might be expected that 
here, as among the Marsupials, where a like condition of the pre- 
caval exists, there would be traces of both right and left azygos. 
And this is precisely what is found among the Rodentia. 
Tt is for this reason that I treat of the Rodents next to the 

The most remarkable condition of the azygos vein so far as 
concerns the Rodentia is seen in the Beaver. But as I have only 
had the opportunity of dissecting a single example of this Rodent, 
the structure may be abnormal in the individual. There is only 
a single azygos; but the vein, instead of lying upon the right 
side, as is otherwise the case when the vein in question is only 
developed upon one side of the body, lies upon the left side. 
This state of affairs is, however, only an exaggeration of that 
found in Jaculus orientalis. In this Rodent, as I have ascertained 
and shall point out, of the two azygos veins present that of the 
left side is the larger instead of, as in the majority of cases where 
there are two azygos and a discrepancy in size between them, the 
right being the larger. 

This condition of the azygos, however, exists among the Mar- 
supials, as has already been pointed out*, and also among the 
Ungulates +, but apparently nowhere except as an anomaly. 

Of Capromys pilorides I have had the opportunity of studying 
two examples which, although they present certain differences, 
agree in certain main features in which they also present unmis- 
takable points of likeness to their allies Hystrix and Dolichotis. 

The azygos is either only or almost entirely to be found on the 
right side~. The vein on that side in one specimen follows after 

* Supra, p. 202. + Supra, p. 190. 
t Dobson (P. Z. 8. 1884, p. 248) only mentions a left azygos. 


an anterior branch from the right jugular. This latter branch 
divides soon after issuing from the jugular (to reverse for the 
convenience of description the direction of the blood-flow) into 
two, one anterior and the other posterior. Each of these supplies 
several intercostal spaces, the anterior two and the posterior 
three. The azygos proper does not branch until after it has 
passed the region supplied by the branch referred to. Its first 
twig supplies two intercostal spaces. The rest are single branches. 
On the left side of the body is a single vessel supplying only one 
space. In the other specimen I observed no branch on the left 
side of the body. On the right side the first branch of the azygos 
runs forward and supplies on each side of the body three inter- 
costal spaces. It obviously corresponds to the anterior branch of 
the jugular of the last individual. 

In Celogenys paca 1 have observed rather different arrange- 
ments of these veins in two specimens. This animal, unlike 
Dolichotis, has two jugulars. In one individual (both were males) 
the azygos was developed on both sides of the body though con- 
siderably longer on the right. On this side of the body a branch 
is given off early in the course of the azygos which supplies 
several intercostal spaces; the main trunk of the vein runs of 
course parallel with this, and gives off no branches until the last 
of those supplied by the large branch referred to. In this speci- 
men the inferior intercostal veins do not open independently into 
the jugular, but into the azygos before it communicates with the 
jugular. In the second individual both right and left superior 
intercostals were quite independent of the azygos of thei side, 
entering each jugular separately. The azygos vein of the left 
side of the body although present was very slightly developed, 
apparently collecting blood from only one intercostal space. 

In Dolichotis patachonica the conditions of the azygos were as 
follows:—The trunk of the right side is the chief one to be 
developed. It is, however, as in some other forms (e. g. Wacropus 
dorsalis *), derived from two branches which arise separately from 
the jugular. The first is a small branch which runs in a forward 
direction. The second trunk is the main azygos. ‘This at once 
gives off a branch which divides into three twigs, the rest of the 
branches are single in their origin from the azygos. On the left 
side of the body a vein arises three or four ribs further forward. 
Before reaching the parietes it gives off a slender twig which 
runs backwards along the carotid and the aorta. I have not 
traced this vessel out to its end; it is evidently to be compared 
with a similar vessel in Lemur macaco. When it has reached 
the parietes this azygos divides into two vessels which run in the 
same straight line and are continuous fore and aft along the body- 
wall. Two intercostal veins arise from the anterior section and 
three from the posterior section. I did not detect any connection 
of this left-hand vessel with the subclavian vein anteriorly. 

* But in this case it is the left azygos which shows the peculiarity mentioned. 

Proc. Zoot. Soc.—1907, No. XIV. 14 

210 MR. F, E. BEDDARD ON THE [ Feb. 19, 

In another example of Dolichotis patachonica, the conditions 
observed were a little different. The azygos proper was only to 
be found on the right side of the body, in correspondence with the 
fact that this rodent has but one jugular vem. In front of the 
azygos are two separately arising superior intercostal veins. On 
the left side the vein described in the first specimen of Dolichotis 
also existed, but was of much more limited extent than in that 
individual. It corresponded in place of origin with the upper of 
the two separate superior intercostals of the right side. The 
vessel divided into two branches only, of which one was behind 
the point of emergence of the vein from the jugular. It is clear 
that this trunk is the same as that which in the other specimen 
takes up the place of the azygos of the left side of the body, though 
it is much less extensive. There is thus no essential difference 
in respect of the veins of the intercostal system between the two 
individuals. Only a difference of degree in the development of 
the same, and that a slight one. 

The Porcupine (Hystrix cristata), belonging as it does to the 
same subdivision of the Rodents as the Patagonian Cavy, naturally 
shows some resemblances to that rodent in the disposition of the 
azygos veins. The right-hand trunk is the chief one to be 
developed *. In front of it, however, arise from the right jugular 
separately two intercostal veins. The main trunk gives off first 
a branch which supplies one rib. This is followed by a more 
important branch which supplies three intercostal spaces as does. 
the corresponding vessel in Dolichotis. After this the branches 
are single. On the left side of the body the vena cava anterior 
sinistra (the left jugular) gives off two vessels following each other 
which arise at about the same level as the two branches of the 
right jugular which he in front of the azygos proper. The second 
of these is the most important, and runs for some distance down 
the body, giving off four branches to the intercostal spaces. All 
these lie posteriorly : there is no forward continuation of the vein 
such as occurs in Dolichotis; but the forward extension of the 
vein in Dolichotis is evidently replaced in Hystrix by the separate 
twig arising from the left jugular. 

In a second example of Hystria cristata, a male, the arrange- 
ment of the azygos veins (text-fig. 72) was apparently identical. 
T use the word “apparently” because in the first example I did 
not count the ribs between which the various branches of the 
azygos ran. In the second specimen the azygos flowed into the 
right jugular at a point on a level with the interval between 
the third and fourth ribs. The first branch, exactly in the same 
way as in the example I have just described, supplied three 
intercostal spaces, beginning with the third rib. A superior 
intercostal vein arose from the jugular in front of this, and 
received from the first and second ribs a branch each. On the 
left side the azygos proper was very slender, but supplied ribs 

* Parsons (P.Z. S. 1894, p. 685) states that there is “only one azygos vein” in 
Atherura africana. 


four to seven, and there was in the same way a superior inter- 
costal. The constancy of the first branch of the azygos of the 
right side which supplies several branches is remarkable. It 
seems to me to be possible that this branch is the real post- 
cardinal, persistent here to a greater extent than in most other 
mammals which I have dissected. In any case its relations are 
perfectly consistent with such a view of its nature. I may 
mention that in this example I noted an esophageal twig arising 
separately from the left jugular not in common with the azygos. 


ry iL) 

Az. l, 

Azygos veins of Hystrix cristata. 

ces. (Esophageal vein. Other letters as in text-fic. 62. 

In the Canadian Porcupine, Hrethizon dorsatum, the jugular, 
contrary to what is to be found in Hystria, is a single vessel. 
The system of azygos veins is also single and confined to the 
right side of the body. There is no trace that I could discover- 
of either azygos or superior intercostals on the left side. The 


212 MR. F. E. BEDDARD ON THE | Feb. 19, 

azygos presents no features of any special interest ; it is like that 
of many other animals. In front are two separately arising 
superior intercostals. The difference between these veins in the 
present genus and in the true Old World Porcupines is one 
more justification of the absence of near affinity between those 
superficially similar rodents. I examined two specimens of this 

Myopotamus coypu has two jugulars, and there is an azygos 
vein opening into each at the same level. That of the right side 
of the body is the one which is fully developed and runs a long 
way down the body. On the left side a slender twig joins the 
jugular of that side which appears to emerge from one intercostal 
space only. The arrangement of these vessels in the Chinchilla is 
the same. 

In Lepus europeus, Arctomys marmotta, Cynomys ludovicianus, 
Lagomys roylei, there is but one azygos and that on the right side 
of the body. 

In Sciurus bicolor there are also two jugular vems as in the 
species which have just been described. There is also in the same 
way but a single azygos vein arising from the right jugular. The 
branches of this vein are collected from the series of intercostal 
spaces commencing with that lying between ribs 4 and 5. 

Seiurus maximus differs from the other Squirrel mentioned in 
the present communication in that there is but a single jugular 
vein, which is the right-hand one of other Squirrels. Into this 
opens the well-developed right azygos vein, which presents no 
differences from that of other members of the genus Sciwrus, 
except of the smallest detail. Its affluents commence with that 
of intercostal space 5/6. Corresponding to this vein, however, 
there is a very small left azygos. This vein collects blood only 
from ribs 4 and 5, and it opens into the single jugular just a 
trifle in front of the point of opening of the large right azygos. 
The right azygos, I should add, in both the species that have just 
been referred to extends back considerably further than the 
last rib. 

Of Sciurus vulgaris I have examined two individuals, one a 
male and the other a female. The two agreed in every detail of 
structure. This species belongs to that group of the rodents which 
possess two jugulars. But in spite of this the azygos vein is 
single and is present on the right side only. “I did not observe 
any superior intercostal vein arising from the jugular in front of 
the point where the true azygos springs. I found the same to be 
the case with Sciwus palmarum. 

In Hydrocherus capybara there are similarly two jugulars and 
only a single azygos, which as in Seiwrus is on the right side of 
the body. There was nothing to be seen of an azygos on the left 
side, which is perhaps remarkable in view of the relationships of 
this rodent. This statement moreover applies to two examples of 
the species. 

In the Jerboa, Jaculus orientalis, a rodent with two jugulars 


also, there are two azygos veins well developed. I have dissected 
two examples, both females, of this species, and find that in both 
the left azygos is rather the larger of the two, markedly so in one 
specimen. This is a curious reversal of the prevalent arrangement 
in Rodents and is reminiscent of conditions found among the 
Marsupials. So far as my experience goes this condition of the 
azygos veins is unique among the Rodents. It obviously culmi- 
nates in the condition observable in the Beaver, where there is 
but a single azygos vein and that of the left side. Precisely the 
same series is to be met with among the Marsupials. 

(9) The Azygos and other Thoracic Veins in the Young of 
Myopotamus coypu, and the Homologies of the Azygos in Mammals. 

A recent examination of four young specimens of the South 
American Myopotamus coypu has furnished facts of great in- 
terest in connection with the real nature of the azygos veins in 
the Mammalia generally, besides making a contribution to the 
anatomy of these veins in that particular rodent. The four 
individuals were born dead, but apparently were of full time. 
They came into my hands on the following morning, and were in 
admirable condition for study. All four agreed excepting in one 
detail, and this apparent divergence may have been due to the 
difficulty in tracing the smaller branches of veins. In all of these 
examples there were three veins in the thoracic region, as well 
as—of course—the post-caval. Two veins (text-fig. 73) were 
symmetrical with each other, and lay on either side of the ver- 
tebral column near to the point of articulation of the ribs. Of 
these veins the right-hand one was constantly the larger. Both 
opened into the right and left anterior cava respectively. Branches 
arising intercostally were seen in both cases. In front of the 
point of entrance of this vein into the precava there were no 
representatives of the venz intercostales supreme. That is to say, 
no veins to which this term might be applied opened separately 
into the precava. But as a matter of fact all the intercostal 
spaces from the very first backwards had their intercostal veins, 
which in the case of the anterior ones passed backwards to join 
the main trunk. They joined the main trunk very nearly at its 
point of entrance into the precaval vein, and the slightest shifting 
of the posterior set of ees would civide oe affluents of 
the precaval into two 
vena intercostalis suprema, anal a Seon azygos. But is this 
vein to be considered an azygos? This point may be deferred 
until after description of the third thoracic vein. This vein was 
rather larger than either of the paired veins just considered. 
It lay in each case on the right side, or—to speak more accurately 
—it poured its contents into the precava on the right and not 
into the left-hand one of those veins, paired here as in many 
other Rodents. The vein really lay rather medianly in position, 
running over the centra of the vertebre. Anter iorly it jomed 

214 MR. F, E. BEDDARD ON THE [Feb. 19, 

the vein of its side which has just been described and opened 
in common with it into the precava. 

In one at any rate of the Rodents this vein bent distinctly to 
the right before effecting this connection, and in another there 
was the same bend and (see text-fig. 73) a slender straight vessel 
continued on the direction of the vein and apparently opened into 
the paired vein rather nearer to the precava. It was generally 
to be seen—possibly invariably present though hard to see owing 
to deficiencies in the blood—that this vein was connected by cross 
vessels with both of the paired vessels already described as running 

Text-fig. 73. 

Lateral view of the thoracic region of newly-born Iyopotamus coypu, dissected 
to show azygos vein (Az.7.) and persistent postcardinal (post.c.). 

alongside of the vertebral column on either side. J am inclined 
to think that these cross trunks were segmentally arranged in 
accordance with the vertebra. This vessel was longer than either 
of the paired trunks, and posteriorly the last cross anastomosis 
that I observed ‘on the right side was continuous with a longi- 
tudinal vessel obviously continuing on the paired vessel of its 
side, though a break occurred between the two. This trunk, 
receiving affluents in the lumbar region, ended by opening into 
the postcava near to the entry into the same of the right renal 
vein. Anteriorly the unpaired vessel lay outside the area which 
would be occupied by the intercostal arteries (though I did not 


actually observe the arteries here); later on it could be distinctly 
seen to lie within that area. The intercostal arteries could be 
observed to lie outside of the vein. We can now consider what is 
the nature of these several veins. There seem to me to be only 
three possible views, one of which is so improbable that it may be 
shortly considered first ot all and then dismissed. This view is 
that the paired vessels are two azygos veins which have been 
formed in a way not indicated in the specimens from the post- 
cardinals, and that the single more median vein represents the 
esophageal branch which is at least very commonly developed. 
from the azygos and often leaves it near to its connection with the 
precaval. If this is the case the vessel must degenerate very much 
in maturity, and its numerous CYross connections with the paired 
veins are not easy to understand. It seems in fact unlikely that 
such unimportant vessels as the visceral branches of the azygos 
should have so portentous a beginning and so impotent a conclu- 
sion, The other alternatives are mutually exclusive. Hither the 
paired veins are the posteardinals and the unpaired vein is a single 
azygos, or the single vein is the one remaining postcardinal and 
the paired veins are azygos veins which have been derived from 
that and another vanished postcardinal. The first of these views 
seems to me to be fairly obviously the correct one; and for the 
following reasons. In the first place, paired postcardinals and an 
unpaired azygos are more likely than the reverse. This is, of 
course, not conclusive, for one ev hypothesi postcardinal might 
have disappeared. Secondly, the presumed azygos corresponds 
with the adult WMyopotamus where there is only one azygos, and, 
as is so general with the azygos, it lies partly outside and partly 
inside the intercostal arteries. The median position too of the 
presumed azygos is like that of undoubted azygos veins, while the 
lateral position of the presumed postcardinals is again In consonance 
with that view of their nature. In the next place, Dr. McClure * 
has pointed out that the azygos arises from the postcardinal in the 
Opossum to the median side of that vein, which is precisely the 
position occupied by the presumed azygos in the young Myopotanus. 
Furthermore, the break between the thoracic and the abdominal 
regions of the paired veins is in accord with the view that they 
really are the posteardinais ; for such a break has been described 
<n the course of the development of those veins. 

It is obvious that these facts throw some light upon the azygos 
veins in general. It seems highly probable that we shall have to 
distinguish in future between real azygos veins or vein and 
persistent posteardinals. It is even possible that the true azygos 
vein is always a single vein, and that when there are apparently 
two-—one on each side—it is a case either of both postcardinals 
persisting or of a single azygos with one persistent postcardinal. 
That there should be such large differences even in allied mammals 
seems surprising; but it is by no means impossible even on 

* Toe. cit. (see p. 183). 

216 MR. F. E. BEDDARD ON THE [ Feb. 19, 

a priori grounds, if—that is to say—we are permitted to use the 
analogy of the development of the postcaval. It is perfectly clear 
from developmental facts that in the Marsupials generally that 
vein has a different origin posteriorly from that which it has. 
in the Rabbit, while McClure appears to have shown that in 
different individuals of Didelphys marsupialis the extreme end of 
that vein originates differently. In man, according to the facts 
collected by Hochstetter, the postcaval has as a variation a 
different mode of origin from that which is normal. 

In the young pouch-living individual of Macropus derbianus de- 
scribed above *, it was pointed out that there were two azygos veins 
in both foetus and parent. There was no trace in either that I noted 
of any other vein. It seems to me to be possible that in that and 
other Marsupials there are not any azygos veins at all, but the 
persistent cardinals take their place. That this is not the case, 
however, in Didelphys appears to be shown by McClure’s investi- 
gationst. Again, the relations of the azygos in Cervus sika 
described abovet lead one to the inference that the anterior 
branch of the azygos running for some distance to the outside of 
the azygos is a remnant of the postcardinal of that side, since the 
junction of the two veins near to their entrance into the jugular 
recalls exactly the conditions which have been described above in 
the young Myopotamus. These remarks, however, cannot be 
considered at present as more than suggested possibilities. It 
seems to me to be equally likely that in those cases where a well- 
developed right azygos or left is accompanied by a less developed 
vein on the opposite side of the body which I have called azygos 
in the preceding pages, the latter is really a persistent post- 

In addition to these there are cases where on one side of the 
body at any rate both azygos and postcardinal appear to persist. 
Those instances present as a permanent condition the state of 
affairs which occurs temporarily in Myopotamus coypu. In one of 
two Celogenys paca described above, there is, as I regard it, a 
well-developed persistent postcardinal on the right side and traces 
of the same on the left, in addition to a well-developed ALY OS 
on the right and a good but not so well-developed azygos on 
the left. In Capromys pilorides there are also considerable 
remains of the right postcardinal as well as of the right azygos. 
In Hystrix cristata (see text-fig. 72, p. 211) we see precisely the 
same thing on the right side; in this species in both examples 
dissected. Dolichotis patachonica has also, if my views are correct, 
both a right azygos and a short right postcardinal. Nor is this 
retention of a feetal condition peculiar to Rodents, though clearly, 
so far as my experience goes, most abundant in them; for it also 
occurs on the right side of the body in Cervus sika. 

The relationship of the posterior end of the azygos to the 
postcardinal of the right side requires further emphasis. As has 

* Supra, p. 202. + Loe. cit. p. 183. t Supra, p. 192. 


been mentioned, the azygos bends over the vertebral column and 
joins a vein running nearer to the ventral side of the body, which 
is hidden posteriorly by the kidney and opens into the post- 
caval near to the entry thereinto of the renal vein. Previously 
to this the vein has collected tributaries from the lumbar body- 
wall. This is precisely what occurs in other and adult mammals. 
It is frequently the case that the azygos, after emitting the vem 
to the last intercostal space, curves in to the same way over the 
vertebral column and can be traced back to the kidney, in the 
neighbourhood of which it debouches into the postcaval vein. 
It seems to be clear, from the relations of the different sections of 
the veins concerned, that the bend to the right (of the right 
azygos) is the equivalent of one of the cross branches, by many 
of which the azygos is put into communication with the right 
postcardinal. Thus the complete azygos in such mammals is 
azygos plus one cross communication with the postcardinal plus 
the lumbar section of the postcardinal. The azygos, therefore, 
has as complex a formation as has the postcaval itself. 

This condition of the azygos in many mammals contrasts with 
the conditions to be observed among the Marsupials. Here, as 
has been pointed out, it is not infrequent for one azygos, either 
right or left, to be a vein of very considerable importance, not 
merely in the thoracic region but also in the abdominal region. 
In various Diprotodont Marsupials, duly recorded in the pre- 
ceding pages, I have found a thick vein in the kidney region, 
sometimes even extending further back, in fact quite into the 
pelvic region. This vein, which in the abdominal region runs 
parallel with the posteaval, is connected with that vem by an 
anastomosis. Further forwards it is directly continuous with the 
azygos vein of its side, which lies in or very nearly in the same 
straight line with it. Not unfrequently there is a thinning 
before the two veins meet, but still a continuity. This was seen 
for example in a specimen of -Zpyprymnus rufescens. In others 
there was no such thinning. These cases are, as it appears to 
me, to be best explained on the assumption that here the “azygos ” 
of the adult is in reality the persistent postcardinal of its side. 
Hence the absence of the bend such as occurs in iJ/yopotamus 
coypu. The thinning I put down to the “break” which has 
already been referred to as occurring between the thoracic and 
abdominal parts of the postcardinal in many mammals 

There are two other matters for consideration in view of this 
suggestion with reference to the differing nature of the azygos in 
the adult. Firstly, the variability of these veins becomes of a 
different aspect, since it is not one but two distinct veins or pairs 
of veins which vary. The actual variability is therefore reducible 
in consequence, and to be considered from different points of 
view. The variability in the proportions of the two azygos veins 
in the Diprotodont Marsupials is, if my suggestions be correct, 
obviously not the same thing as the variability in the two veins 
which bear the same name among the hollow-horned Ruminants. 

218 MR. F. E, BEDDARD ON THE [ Feb. 19, 

Broadly speaking, it appears to me that the true azygos is by no 
means so variable a vein as are the persistent postcardinals, a 
conclusion which is in perfect accord with general principles. A 
partially persistent, though obviously decaying structure, is apt 
to vary more than a newer structure, which may be increasing 
rather than diminishing in importance. Thus we find that 
the true right azygos 1s very constant among the Carnivora. 
Secondly, there is the fact of the continuation into the abdo- 
minal region even as far as the pelvic region of the azygos vein. 
In some cases at any rate, possibly in the majority or even all 
those animals which (e. g. Carnivora) possess a true azygos vein, 
i. €. secondary vein not the persistent postcardinal, that vein is 
continuous posteriorly with a longitudinal vein which receives 
lumbar branches and opens indirectly or directly into the post- 
caval vein. This part of it I regard, for reasons already put 
forward, as a portion of the persistent postcardinal. Now it is 
precisely among the Diprotodont Marsupials, whose azygos veins 
are, as I think, true postcardinals, that we find a very large vein 
continuing into the ebdominal and even pelvic region of the 
body *. It seems to me that there is some significance in this 
fact, and that it tends to support my contention that among 
adult mammals the azygos veins fall into two categories. 

(10) The Condition of the Azygos and the Classification 
of Mammals. 

It is clear from the facts here set forth, that the condition of 
the azygos vein or veins has a distinct relation to the mutual 
affinities of the several groups of Mammalia, but. in quite a 
general way. ‘Thus the Lemurs agree with the other Primates in 
having only the single right azygos ; and, moreover, it is in these 
two groups alone that an hemiazygos is at all prevalent on the 
left side, joining the azygos some way before the latter opens into 
the right vena cava superior. All the members of the Order 
Carnivora agree with each other in the same character; the. 
divergences in these three orders or two orders being of the 
slightest when compared with some other groups. I do not 
infer from this that there is a special relationship between the 
Carnivora and the Primates of course. Both groups have, as I 
think, independently arrived at the same state of affairs as 
regards the azygos vein. But for each group considered alone 
the facts at least agree with the general view of their affinities. 
It is important to note that the Lemurs agree with the Apes, 
and the Arctoid Carnivora with the Ailuroid. No difference in 
mode of life seems to have affected this deep-seated character. 
The Otter cannot be distinguished in this particular from the 
Civet Cat, or the Raccoon from the Suricate. 

* Hochstetter, however, declines to recognise as persistent sections cf post- 
cardinal Robinson’s description and figures of a left postcaval extension of azygos 

(Studies in Anat. Owens Coll. i. 1891, pl. vi. figs. 1, 2, &c.). But it does not 
follow that this objection would apply to Macropus. 


Again, the cavicorn Ruminants mostly agree with the Suidz in 
the entrance of the left azygos into the heart independently of 
the remains (if they exist) of the left vena cava anterior, And in 
this Moschus and Hydropotes agree with them. This agreement 
is not without interest, particular ly when it 1s considered that the 
Cervidee may differ in the relations of the azygos: in Cervus sika 
the right azygos being the prevalent or only azygos and entering 
in the! usual way the right anterior cava. Most zoologists sepa- 
rate the solid-horned from the hollow-horned Ruminants, though 
to others the distinction between the two groups of Artiodactyles 
appears “fanciful.” In any case the Perissodactyles have their 
own plan of azvgos structure, which happens to agree with that 
of some Cervide, but distinguishes the Suborder from the Pigs 
and hollow-horned Ruminants. In connection with the classifi- 
cation of the Ungulates, as supported by the disposition of the 
azygos veins, the position and number of these veins in Hyrax 
are of particular interest. In this ‘“‘subungulate,” admittedly 
primitive, and standing nearer to the base of the Ungulate series 
than any other living form, except the Elephant, there is as it 
were a hesitation to adopt definitely the form of the azygos veins 
to be seen in either Artiodactyle or Perissodactyle. 

It may also be fairly said that a low position in the series is 
indicated by the persistence of both azygos veins. These persist 
in Hehidna, and are very generally prevalent in the Marsupials, 
the Rodentia, and, though here our knowledge is more deficient, 
in the Insectivora. It is the general opinion that of these groups 
at least three are primitive groups, and many would think the 
same of the Rodentia. The division of the Marsupials into 
Diprotodont and Polyprotodont is justified by the condition of 
the azygos veins; for in the Diprotodont division there is a much 
greater tendency for the two azygos veins to persist than among 
the Polyprotodonts. Furthermore, as emphasising the gap that 
separates the Marsupials from most of the higher mammals, it is 
noteworthy that in them (7. e. the Marsupials) it is the left azygos 
which is apt to be predominant and not the right vessel. In the 
Order Rodentia also the conditions of the azygos agree with 
current views as to their subdivisions. On the whole the 
Hystricomorphine Rodents are those im which the two azygos 
veins persist to a greater or less extent. An exception to this 
statement would appear to be the genus Dipus. But it must be 
remembered that the late Dr. Dobson concluded from certain 
facts in muscular anatomy that Dipus was not remote from the 
Hystricomorpha *. 

(11) The Position of the Azygos with reference to the Vertebre. 

In dealing with the development of these veins in Mammals, 
Hochstetter has dwelt upon the fact that the heart moves back- 
wards and with it of course the blood-vessels, including the 

* Journ. Anat. Phys. xvii. 1883, p. 177. 

220 MR. F. E. BEDDARD ON THE [Feb. 19, 

Ductus Cuvieri, attached thereto. The opening of the posterior 
cardinal veins into the Ductus Cuvieri thus undergoes an altera- 
tion of position with reference to the thoracic vertebre. Thus 
Hochstetter found that in the embryo of a rabbit fifteen days 
old “ Der Stamm der V. azygos miindet schon in der Hohe des 
Zweiten Intercostalraumes in die Hohlvene ihre Seite,’* while 
in the adult its position is further back. It is interesting to note 
in relation to such developmental facts the varying position of the 
- entrance of the azygos into the precava in different families and 
genera of adult Mammals. It would seem plain that if the 
azygos vein or veins enter the precaval further forward in one form 
than in another, the conditions are in that type more primitive. 

On the whole it would appear that as a general rule the point 
of entrance of the azygos vein is opposite or about opposite to 
the fifth rib. It is at any rate so in Macropus, Lutra, Sciurus, 
Suricata, Equus, Lemur, &e. It seems to be the general position 
among the Carnivora and Primates. I have not met with any 
instance of a shifting backwards further than this point, more 
than the very smallest. I have noticed occasionally that the 
sixth rib instead of the fifth is the first to emit an intercostal to 
the azygos. This was the case, for example, with a new-born Ovis 
tragelaphus. Now, although Hochstetter and McClure (working 
it is true with different types) disagree as to how much exactly 
of the embryonic postcardinals persist in the adult as the azygos 
vein or veins, there is no doubt that the actual orifice of the 
azygos into the precava represents the embouchure of the post- 
cardinal into the Ductus Cuvierl. We can, therefore, argue 
concerning the shifting position of this point. Dr. McClure has 
himself observed + that the left azygos of Didelphys ‘“ opens into 
the left precava about opposite the head of the third rib,” while 
“the right azygos vein, when present in the adult, opens into 
the precava about opposite the head of the second rib.” It cannot 
be said, however, that this position of the azygos, which is to be 
regarded as a more primitive position, is In any way characteristic 
of the Marsupials. It does nevertheless occur in that group, and 
1 have found the same in Psewdochirus peregrinus. It is in- 
teresting in relation to this matter to find that in the Insectivore 
Erinaceus the Ductus Cuvieri on both sides of the body les 
opposite to the interval between the second and third ribs. The 
Rodents also show to some extent the same anterior position of 
the point of entrance of the azygos into the precaval. Thus in 
Hystrix cristata the first affluent of the right azygos arises between 
the third and fourth ribs. On the other hand, among the Car- 
nivora and Primates, as already said, this forward position of the 
entrance of the azygos is never (in my experience) to be met 

Indeed, the facts stated at length in the preceding pages, which 
have just been briefly reviewed, point distinctly to the persistence 

* Morph. Jahrb. xx. 1893, p. 573. 
+ American Journ. Anat. 1906, p. 185. 


of a more primitive position of the Ductus Cuvieri in animals 
which possess as a rule two azygos veins, and are thus also 
primitive in this peculiarity, and which moreover are usually 
assigned on these grounds to a low place in the Mammalian 
series. The whole group of facts fit in with each other in perfect 
harmony. J believe that these facts are not without importance 
in considering the Artiodactyle Ungulates, or rather the hollow- 
horned Ruminants and the Pig tribe, which agree in the characters 
of the azygos veins. 

The right azygos in these animals is always situated far forwards, 
beginning even (e. g. in Oryx beatriz) with the second intercostal 
space, and although the left and prevalent azygos does not occupy 
what is ea hypothesi a primitive position, there are nevertheless 
true azygos veins present. The anterior section of the azygos, 
which runs forward of the point of opening into the heart, is 
perhaps to be regarded as the left precaval otherwise missing. 
There is thus some agreement with the suggestions already 

(12) The Venw Intercostales Supreme. 

These veins, so named by Hochstetter, and often termed inferio1: 
intercostals, are superficial veins made up of one or more of the 
intercostals anterior to those which unite to form the azygos 
vein or veins. ‘They are the superficial superior intercostal veins 
of McClure*. As will be gathered from the foregoing pages, these 
veins are not invariably pr resent as trunks opening into the vena 
cava superior; nor indeed are they invariably present at all. I 
presume that in these cases, where no such veins are apparent, 
the circulation is effected by ‘the deep series of intercostal veins 
lately figured with great elaboration in the Opposum by Dr. 
McClure in the paper already referred to. The details of the 
occurrence of these veins in various genera of mammals have been 
already dealt with in the foregoing pages. From these it will 
appear that the veins in question are by no means of universal 
occurrence, and that when they do occur they are not always 
connected with the precaval vein or veins. In man, for instance, 
these veins are described as entering the left and right vene 
anonyme respectively. This appears to hold good for the Primates 

But I do not profess in the present communication to have dealt 
at any length with the Primates. Among the Carnivora these 
veins are evidently not common, and when they do occur some- 
times join the precava and sometimes the vena or ven anonyme. 
In the Marsupials also the venze intercostales supreme are not 
universal, but when they are met with they seem as a rule to 
join the precaval or precavals. These veins are much more 
general among the Hystricitorm Rodents, where indeed, so far 
as my observations go, they are practically universal and join the 
precaval. As a rule, too, in this group the veins are paired and 

* Amer. Journ. Anat. vol. 11. 1903, p. 371. + Loe. cit. fig. p. 380. 



each precaval has its own separately debouching intercostals. 
That this is not at all due to the prevalence among these Rodents 
of two precayals is shown by the same constancy of the ven 
intercostales supreme among the Artiodactyle Ungulates. The 
exceptions that I have met with to the rule that these veins are 
present on both sides and open into the single precaval are but 
few in number. Concerning other groups the facts at my dis- 
posal are so few that I hesitate to attempt any remarks of a 
general character. 

(13) Conclusions. 

Although the facts detailed in the preceding pages, as well as 
those made known by the records of others, might be largely 
added to, enough appears to me to be known to permit of a few 
general observations upon the azygos vein in the Mammalia. 

(1) With the exception of the Cetacea the azygos is always a 
well-developed vessel of fair size among the Mammalia, and 
generally to be found fully developed upon one side of the body 
only, and this the right. Its functions appear never to be taken 
up by the vena cava inferior, so that 1t is never rudimentary. 

(2) There is only a right azygos with no traces of a left— 
except rarely as a hemiazygos not reaching the vena cava anterior 
—in Carnivora, Lemurs, American Edentates, and Primates. 

(3) The existence of two azygos veins characterises several 
Marsupials, Rodents, Insectivora, and most Artiodactyles. In the 
two first-named groups the existence of the two veins is associated 
with the presence of two superior vene cave. It is not, however, 
every Rodent with two superior cavee which has also two vene 
azygos. But, as it appears, no Rodent with one superior cava 
only has more than one vena azygos. The Artiodactyles are 
exceptional in that, while possessing only a single anterior vena 
cava, there are two azygos veins. But the left azygos generally 
opens into the right auricle with or close to the vena cava posterior. 
Rarely there is also a connection with the single vena cava 
anterior. Where two azygos veins are present it is unusual for 
them to be equally developed. Hither the right or the left is 
well developed and the other smaller or much reduced. 

(4) The existence of a left azygos only characterises a few 
forms belonging to the groups Marsupials, Rodents, and Artio- 
dactyles. This condition of the azygos is not, like the two which 
have been dealt with, characteristic of any large group. 

(5) It is to be remarked that, on the whole, the possession of 
two azygos veins is associated with a low position. 

(6) With certain exceptions (e. g. Hpyprymnus rufescens) the 
azygos vein is fairly constant in its characteristics for a given 
species. This is sometimes very markedly the case. 

(7) As will be gathered from the details given, the azygos is 
of some use in the classification of Mammals. Thus Hyrax shows 
its Ungulate affinities, and several genera of Hystriciform Rodents 
agree together in the character of the azygos veins, &e. 


(8) As a general rule the entrance of the azygos vein or veins 
into the precaval or precavals is more anterior in position in 
Mammals, occupying a lower position in the series than in more 
specialised types, 7. €. opposite to second rib instead of fifth or 
sixth. This corresponds with ontogenetic shifting back of heart 
and blood-vessels. 

(9) The conditions observable in the newly-born young of 
Myopotamus coypu seem to show that the posteardinals may 
persist as such at least up to the time of birth, and in some adult 
Rodents one is also persistent. 

(10) The same species shows that the azygos of the adult is 
independent—except for a very short tract at its opening into the 
precava—of the postcardinal of its side, thus confirming the embryo- 
logical results of others who have affirmed that only the very 
commencement of the azygos is traceable to the persistent post- 
cardinal of its side. 

(11) It is probable that the veins called “azygos” in adult 
Mammals are not in every case strictly homologous veins. Where 
there is but one azygos present (e. gy. Carnivora) it is probable 
that that vein is the true azygos, except in the abdominal region 
where it is formed by the persistent postcardinal. In cases where 
there are two azygos veins both may be (? certain Marsupials) 
persistent postcardinals, or one of the two may be a remnant of 
the posteardinal, the other being a true azygos. 

(12) These and some other facts and conclusions lead to the 
inference that the true azygos vein of Mammals (7. ¢. that formed 
by an outgrowth of the postcardinal) is a structure which has been 
developed in the Eutheria. 

5. Ideas on the Origin of Flight. 
By Dr. Baron Francis Nopcsa. 

[Received February 8, 1907.] 
(Text-figures 74-82.) 

Although much has been written on the origin of flight, 
yet till now no really satisfactory explanation for this kind of 
locomotion has been found. ‘This is, so far as I can under- 
stand, mainly due to the fact that it has on @ priori grounds 
been supposed that all the principal groups of flying vertebrates— 
namely, Pterosaurs, Bats, and Birds—originated in a similar 
manner, without fully appreciating the fundamental fact that, 
from the mechanical standpoint, patagium and feather are two 
perfectly different organs. 

A patagium is a soft flexible membrane and in consequence 
requires, to be effective, numerous firm radial supports originating 
from the body that has to be carried, whereas for a series of 
semirigid but elastic quills one line of attachment is sufficient. 

In consequence of this difference, a patagium-flyer must always 

224 BARON NOPCSA ON THE [ Feb. 19, 

adapt fore and hind limbs and tail to the support of the patagium, 
whereas in a generalised feathered animal only the feather- 
supporting elements need become affected by violent specialisation. 
The development of the posterior limb in such an animal is but 
little, if at all, affected by the development of flight (foot in Eagle, 
Parrot, Woodpecker, Nightingale, Goose, Stork, Ostrich, &c.). 

As to flight itself we have to distinguish, as partially already 
pointed out by Dollo, three distinct stages of evolution : first 
parachute or passive flight, then flight by flapping the wings or 
Night by force, and lastly soaring or flight by skill. 

As Langley and Lucas pointed out in their highly interesting 
papers, the soaring birds lack carrying power (in accordance with 
which fact the crista sterni is often comparatively feebly 
developed), while flight by flapping of the wings, as shown by 
the generally soaring Eagle when carrying prey, enables the 
animal to support a good deal of weight. 

That soaring birds show a sharply pointed wing, while birds 
that fly mainly by flapping display a wing with a more or less 
rounded outline, is well known. 

After these preliminary, but I think essential, observations, 
T shall now point out some characters of Pterosaurs, flying 
Mammals, Dinosaurs, and Birds that have not yet been brought 


The Dimorphodon, till now the -earliest-described long-tailed 

Text-fig. 74. 

Hind limb of Dimorphodon 

Pterosaur, shows ‘in its hind limb no sign whatever of cursorial 
locomotion (text-fig.74). The metatarsals 1-4 are equally developed, 

1907. | ORIGIN OF FLIGHT. 225 

but the 5th is somewhat thicker and also much shorter. The 
elongate phalanges of the 5th toe further prove clearly that no 
cursorial adaptation modified the form of these bones. Very 
much the same type of foot is visible in the equally long-tailed 
Campylognathus; and when we turn to the Rhamphorhynchus of 
the Solenhofen Slate (text-fig. 75), we find not only no cursorial 
modification of the four-toed slender foot but quite decided degene- 
ration. However, according to Zittel, the number of phalanges in 
the 5th toe is perhaps somewhat greater than in the drawings 
given for Dimorphodon or Campylognathus. Since the spur-like 
clawless 5th digit of the foot is very strongly developed in Dimor- 
phodon, there is, as Owen observed, good reason to believe that a 

Text-fig. 75. 

Hind limbs of Rhamphorhynchus. 

uropatagium was not only present but even very well developed ; 
whereas we know that in Rhamphorhynchus, im accordance with 
the less developed 5th toe, no uropatagium extended to this part of 
the body. The resemblance of the Rhamphorhynchus sternum 
to that of the Bat (Z’aphozous) likewise has to be noticed. 

That in the Liassie Dimorphodon the wing-finger is relatively 
shorter than in the Tithonian Rhamphorhynchus is a fact so- 
obvious as scarcely to demand attention. The short-tailed 
Pterosauria of the genus Pterodactylus, with comparatively short 
wing-bones, resemble Campylognathus in having four feeble and 

Proc. Zoo. Soc,— 1907, No, XV. 15 

226 BARON NOPCSA ON THE [Feb. 19, 

equally strong metatarsals which all approximately attain the same 
length, while, in harmony with the lack of a tail, of the 5th toe 
only a rudiment (text-fig. 76) now remains. Among Chiroptera we 
find that in the tailless forms, notably Péeropus, quite similarly 
the os calear of the hind leg is less developed than in the long- 
tailed species. 

Hind limb of Péerodactylus. 

(The rudiment of the 5th toe is unfortunately not shown in this drawing.) 

On account of the anterior prolongation of the ilium in 
Pterosaurs, and on account of the great number of vertebre united 
in the sacrum, it has been frequently assumed that the Pterosaurs 
enjoyed a bipedal locomotion. Both these arguments, however, 
fail to convince me, and this principally on account of Vyctosaurus, 
which, although certainly not a bipedal genus, has a still greater 
number of sacral vertebre, and because in Pteropus there is like- 
wise a pseudosacrum present. Another argument that can be 
brought forward as annulling the hypothesis just mentioned 
consists in the fact that the Pterosaur pelvis, though showing 
considerable length, has an ilium of an exceedingly low and 
narrow Bat-like outline. 

A Pterosaur of whose crawling habits we can be quite sure is, 
as just mentioned, the Upper Cretaceous Vyctodactylus, for, as 
Williston pointed out, the acetabulum is placed far back, nearly over 
the edge of the sacrum, so that it was impossible for the knees in this 
animal to meet in the middle, and at times the knees may even 
have been turned more or less backward. When the femora 
were rotated outward and abducted, the tibie might have been 
brought parallel with each other. Exactly similar conditions are 

1907. | ORIGIN OF FLIGHT. 227 

to be met with among Bats, whose crawling locomotion is familiar 
to every student. 

The elongation and attenuation of the hind feet in Vyctodactylus 
(text-fig. 77) are also characters that demand mention, and a 
similar elongation is again to be met with in the tailless Vampires. 

An interesting feature is the co-ossification of numerous dorsals 
in Vyctodactylus and the nearly allied European genus Ornitho- 

Text-fig. 77. 

Hind limb of Nyctodactylus. 
(Photograph of specimen in the British Museum.) 

A Triassic long-tailed Pterosaurian, Zribelesodon (the detailed 
description of which I intend publishing on some other occasion), 
shows much the same proportion between total length of hind 
and fore limbs as does Galeopithecus ; and, although Galeopithecus 
proves to be in no way related to Chiroptera, still we must suppose 
that the whole order Chiroptera, considering the patagium, passed 
through a Péeromys- and a Galeopithecus-like stage in the course of 
its evolution. 

Since, as already pointed out, a patagium requires many spear- 
like supports, and since in arboreal animals fore and hind limbs 
are to the same extent used for running and leaping, it is evident 
that primarily fore and hind limbs must have become to the same 
extent used for the support of the patagium, which necessarily had 
to take origin at the centre of gravity between humerus and femur. 

As soon as such a potentially flying animal became actively 

228 BARON NOPCSA ON THE [ Feb. 19, 

volant, and began to fly by force—that is, move its patagium, 
—the sternum must evidently have developed a sternal crest, 
the patagium must have continued to increase its surface, and 
this then would not only produce stretching of the limbs but also 
development of secondary supports of the patagium. 

Such secondary supports are, as we shall see, developed at 
different times and in different ways, being produced by special 
development of the olecranon, the carpal bones, and _ossified 
tendons. In the long-tailed Pterosaurs such supports are absent 
in the fore limb; in the short-tailed Pterosaurs, however, they are 
well developed and are represented by a modified carpal which, 
according to Williston, shows (Pterodactylus — Nyctodactylus) 
progressive evolution. Another modification that each patagium 
produces in the animal’s body is to bring all the radial supports 
to the same level, and this, making the acetabulum and knee 
rotate outward and backward, produces subsequently crawling 

When the last stage of development is attained and aerial 
locomotion accomplished by skill and not by force (Wyctodactylus, 
Rhamphorhynchus), the patagium obviously not only would assume 
a pointed outline and become reduced to a smaller surface, but in 
some cases also the tail would change to a rudder-like organ 
(Rhamphorhynchus) or become entirely lost, while the attenuated 
feet would in this case assume the function of steering (Vycto- 
saurus). It is of no small importance that of the two highly 
specialised groups of Pterosaurians(Rhamphorhynchide and Nycto- 
sauride) the tailless ones should have survived the longer. 


Very much the same changes as are to be found between 
Tribelesodon, Dimorphodon, and khamphorhynchus on the one 
hand, and Péerodactylus and Nyctodactylus on the other, are also 
to be observed when we come to consider the patagium-flying 
Mammals. A set of good diagrams of flying mammals has recently 
been published by R. 8. Lull. Petawrus and all other animals 
with a small patagium represent the stage where, as in all arboreal 
animals, a very long tail is present. 

Only a plagiopatagium is present in Petawrus, a propatagiam 
is added in Péeromys, whilst in Anomalurus even a uropatagium 
is present. Asin Pterosaurs, supplementary patagial supports are 
frequently developed. In Anomalurus and Vespertilio such a 
support arises from the olecranon, in Pteromys it is partially 
attached to the pisiforme and partially, though to a less extent, 
to the 5th metacarpal; and in embryos of Chiroptera quite a 
similar structure is met with: a modification recalling the back- 
wardly directed toe of the hind leg in Dimorphodon is produced 
by the development of the caleaneum’s calear projection. As in 
Pterosaurs, so also in flying Mammals a very low ilium is present, 
and this not only in: Chiroptera but also in Galeopithecus, where, 

1907. ] ORIGIN OF FLIGHT. 229 
even during ontogenetical evolution, a backward rotation of the 
ischium, and in consequence a flattening of the pelvic girdle, is 
to be met with. Pteromys and Anomalurus, according to Dollo, 
have to be termed passive flyers: the first partially active fiyer 
seems to be Gialeopithecus, for, according to Wallace, this animal 
is not only capable of sailing downward, but at the end of its 
downward oblique glide to rise a little upward. 

Galeopithecus, however, is a long-tailed, comparatively short- 
armed patagial animal, in which never theless the patagium ex- 
tends even to the tips of the digits and to the end of the tail ; 
while when we turn from this 0) the specialised actively flying 
Chiroptera, we are impressed firstly by the elongation of the 
wing, and secondly by the frequent partial or total loss of the 

Both in Pterosaurs and Bats the main movement during flight 
seems to have been, and still is, dependent on the humeral articu- 
lation. The similarity of ‘the patagial structure in Lhampho- 
rhynchus and Bats, as remarked by Zittel, is also to be noted. 
The hairless condition of the patagium in ‘Chiroptera compared 
with Galeopithecus is likewise a more specialised feature ; while 
Pteropus vula SG (more specialised than any Bat in regard to the 
caudal region, ‘‘chevauchement de spécialisation ”) * shows, by 
possessing some hair on the interior surface of the patagial 
membrane, an intermediate stage. 

In Pterosaurs, as also in Péeropus, the number of sacrals is 
augmented, and in the latter they even form, by co-ossifying with 
the ischium, a pseudosacrum. 

The more or less perfect reduction of tibia and ulna is another 
character that is noteworthy in all patagium-bearing Mammals. 

in an analogous manner to the Cretaceous Ornithocheirus, also 
in some Bats a rigid thorax is attained, though in this case the 
ribs and not the vertebrze co-ossify. 

Since we may safely assume that Bats descended from Mammals 
which possessed a well-developed neural spine, the reduction of 
this process, noticeable also in the Flying Lizard (Draco volans), 
has also to be considered as a sign of specialisation 

The thin and dense skull-bones also unite in specialised Bats, 
very much as in Birds and Pterosaurs; and as to the brain, 
there exists a great amount of resemblance between the cast of 
the brain-cavity in some Kocene Bats, in Hesperornis, and in 

Only in one point is there a pronounced difference between the 
Pterosaurs and the Bats, and this is in the transformation of the 
phalanges of the pes and manus. While in the Pterosaurs a reduc- 
tion of the claws takes place in the pes, and they remain present 
in the manus, in Chiroptera exactly the opposite happens; but 
this divergence is easily understood when we consider that the 
Chiroptera had, in consequence of adapting four fingers to flight, 
only their hind feet at their disposition, for resting ‘and suspend- 
ing on branches, while the Pterosaurs, which developed only one 

230 BARON NOPCSA ON THE [ Feb. 19, 

wing-finger, could always suspend themselves by the remaining 
free digits. This is, perhaps, also the reason why, in both groups, 
ulna and radius, tibia and fibula have been reduced in a different 
manner. In Birds the same problem has been solved im quite 
another manner (musculus ambiens and peculiarities in the 
structure of the tendon-sheath of musc. flexor. digit. im the 
phalanges). Text-fig. 78 shows the hind limb of a Bat identified 
by Dr. K. Andersen as Hipposiderus gigas. 

Text-fig. 78. 

Hind limb of Hipposiderus. 

Since all flying animals must needs have developed from agile 
quick-moving animals, since all living patagium-flying animals 
(such as Draco volans, and the other living animals mentioned 
in this paper) are arboreal, leaping, quadrupedal creatures, and 
since, further, a bipedal cursorial animal, on account of mechanical 
impossibilities, can never develop a patagium—for such an organ 
would in bipedal (7. ¢., erect) locomotion only catch the air and so 
prevent running without raising the body,—and since the union 
of fore and hind limbs is directly opposed to bipedal cursorial 
locomotion, we can safely state that all patagium-flying animals 
originated from quadrupedal, leaping, arboreal forms. 

Bats and Pterosaurs, though they support the wing in different 
ways, still show an analogous direction of evolution—as shown by 
the development of a patagium with all that this implies; thus we 
may safely state that Bats and Pterosaurs have arisen in similar 
manner from quadrupedal arboreal forms. 

1907. j ORIGIN OF FLIGHT. 231 


In consequence of the quite extraordinary tendency of Dinosaurs 
to specialise every now and then along Avian lines, and in 
consequence of the fact that the most primitive Dinosaurs are 
bipedal in their habits, it is not only probable that all Dinosaurs 
originated from bipedal forms (I only need to quote the numerous 
bipedal tracks in the Red Triassic Sandstone in Connecticut), but 
that they also are very nearly related to the primitive Birds. 

Since Dr. Holland thinks that the Dinosaur likeness to Birds 
is sometimes greatly exaggerated, I would like to mention some of 
the most characteristic primitive and adaptive Avian features of 
Dinosaurian reptiles: basis cranii (Hypsilophodon, Compsognathus), 
development of beak (Orthopoda, beak perhaps developed 
independently in different suborders, caused by latent homoplasy), 
lack of neural spines in cervical vertebrae (Sauropoda), dorsal 
neural spines bifid (Diplodocus), saddle-shaped articulating surface 
of sacrals (Streptospondylus), synsacrum (Orthopoda), 'pyornis- 
like caudals (Diplodocus), Avian scapula (Orthopoda, Theropoda), 
co-ossification with the coracoid (all Dinosaurs), manus (Orni- 
tholestes), ilium covering last ribs (Sauropoda), ilium touching 
neural spines (Stegosauride), ilium showing antitrochanteric 
ridge and dorsal plane (Theropoda) ; backward rotation of pubis 
and subsequent development of processus pseudopectinealis 
(Orthopoda), femur shorter than tibia (many Dinosaurs) ; 
reduction of fourth trochanter (all Dinosaurs), distal end of 
femur (Streptospondylus, &c.), development of processus ascendens 
astragali (Theropoda), fusion of caleaneum and astragalus with 
tibia “(Compsognathus) ; position of hallux (Theropeda) ; pheu- 
maticity or light structure of the whole skeleton (many Dinosaurs). 

In a paper on the evolution of Dinosaurs, I pointed out that 
the Theropoda specialise by developing an interpubic ossification, 
by augmenting the number of their sacrals,; by changing the 
character of their vertebre from biconcave to opisthoccelous, by 
lengthening their neural spines in the dorsal region, and by 
developing a proc. asc. astrag. and reducing the number of their 
toes. In more specialised Theropoda the metatarsals become 
always more closely applied, and. lastly, these animals specialise 
by losing the fourth trochanter. Most of these changes are also 
notable among the bipedal Orthopoda, and since this develop- 
ment is independent of that in the Theropoda, we must consider 
them as homodynamic changes; besides this, in Orthopoda we 
can trace a thickening of the bony matter and the development of 
a processus pseudopectinealis. A functionally analogous osseous 
process is developed in most running birds after the co-ossification 
of the pelvic elements. 

Since we can be sure that in Dinosaurs all the changes 
mentioned are not due to the giving up of volant habits, but are 
merely signs of cursorial adaptation, we have a clue to under- 
stand some of the changes that occur among the Paleognathous 
Birds. Besides this we can fix the fact that the Dinosaurs, like 

232 BARON NOPCSA ON THE [Feb. 19, 

many cursorial Mammals, were not only set the problem of 
developing a flexible dorsal vertebral column, which was attained 
by development of convexo-concave intervertebral articulation, 
but that to this, in consequence of the position of their head, they 
had to add strength in the vertical direction, which could only be 
attained by developing the attachment surfaces for the musculus 
longissimus dorsi on the neural spines and the producing of 
hypapophysis-like knobs on the cervicals. This, moreover, is the 
first consideration adduced since 1887 that shows us that the 
vertebral column of Zgwanodon, though provided with ossified 
tendons, cannot have been altegether rigid. Text-fig. 79 is in- 
tended to show the highly modified foot of the Cretaceous Dinosaur 
Ornithomimus, and can be compared with the feet of Dipws and 

Alactaga (text-fig. 80). 
Text-fig. 79. Text-fig. 80. 


Text-fig. 79.—Hind limb of Ornithomimus. 
80.—Hind limbs of Dipus (left) and Alactaga (right 


Leaving Dinosaurs and turning to Birds, we observe the follow- 
ing salient points :-— 

The first and most primitive Bird we know, Archwopterya 
(text-fig. 81), shows not only a perfectly bird-hke femur and tibia, 
but also tridactylism, and this is, as demonstrated by Dinosaurs 
and the Dipus-like rodents, a prominent feature of bipedal 
cursorial or saltatorial specialisation, while it never occurs among 

arboreal forms. 
The pelvis of Archwopteryx, moreover, is essentially that of a Bird, 

1907. ] ORIGIN OF FLIGHT. 233 

and as a sign of cursorial locomotion there is even an indication 
ef a processus pectinealis. The vertebre are free, and neural 
spines are present. 

Besides this, Archeopteryx differs from all Birds by having a 
long laterally feathered tail, that to a certain degree reminds us 
of the peculiarly covered and flat-looking tails in the mammals 
Acrobates and Ptilocerus low. 

Text-fig. 81. 

eae (i oopkese eee PS 

Hind limb of Archeopteryx. 
(Photograph of the British Museum type specimen.) 

The ossified tendons which occur in the tail of Archeopteryx 
show further that strength of this organ was required just 
as much as in the tail of Dimorphodon or Rhamphorhynchus. 
A long tail, sometimes even with ossified tendons, is quite a marked 
feature of the Dinosaurian bipedal reptiles, and its loss, as shown 
in Pterosaurs, is generallyin harmony with the better adaptation to 
flying locomotion. 

The rounded contour of the Archwopteryx-wing, together with 
the feebly developed sternum, show us that Archwopteryx, though 
perhaps not an altogether badly flying creature, can on no account 
have been a soaring bird, but a bird that was yet in the first 
stage of active flight. 

That the soaring Frigate-Birds and Albatrosses have a com- 
paratively weaker sternum than the Gallinaceous Birds has already 
been mentioned ; and I therefore need only point to the formation 
of a rigid thorax in flying birds as analogous to the condition in 
Pterosaurians and Bats and in opposition to the Ratite, and to 
the fact that the cursorial Paleognathz, contrary to the flying 

Proc. Zoou. Soc.—1907, No. XVI. 16 

234 BARON NOPCSA ON THE [ Feb. 19, 

Tinamous, possessed not only free vertebrve, but even elongate, 
Dinosaurian-like neural spines in the dorsal region, and this 
because also in this case, for running, strength had to be united 
with mobility in the dorsal region, whereas for flight, as already 
mentioned, strength and rigidity seem to be the qualities required, 
go that the neural spines become to a certain extent useless. It 
is especially to the curious dorsal and caudal vertebrae of Apyornis 
that I should like to draw attention. Probably mobility is one 
of the reasons why in the flightless Hesperornis saddle-shaped 
vertebre were developed at a period when Jchthyornis still showed 
biconcave articulation, although I am quite aware that perhaps 
other explanations will have to be sought for, since also in other 
ways Hesperornis indicates a more specialised form, and this not 
only by its wing-bones being already reduced, but by exhibiting a 
certain tendency to lose its teeth, since these are no longer placed 
in distinct sockets as in Archewopteryx and Ichthyornis, but in a 

If we, after these preliminaries, now suppose that Birds, betore 
attaining the Archwopterya-state, originated from quadrupedal 
arboreal animals and only after having learnt to fly became bi- 
pedal, it is difficult to understand why they in general show Dino- 
saurian affinities, why they did not use both hind and fore limbs 
to the same extent for flight as they would have done for arboreal 
locomotion, why the bones of the pectoral region and of the wings 
show more primitive traces than the hind parts of the body, and 
why they did not, like all other quadrupedal flying animals, 
develop a patagium ; whereas, if we consider that in Archwopteryx 
the anterior extremities, though bearing the most important 
ectodermal pinions, are less modified than the posterior extremities, 
which are already perfectly bird-like, and if we then suppose that 
Birds originated from bipedal Dinosaur-like Reptiles, it is easy to 
understand what induced the Birds to attain an Archwopteryx- 
like stage of evolution, for at first a certain amount of bipedal, 
and only afterwards a volant, modification would be required. 

While we can safely state that a bipedal animal never could or 
did develop a patagium without giving up bipedalism, this cannot 
be said of feather-bearing forms, for we may quite well suppose 
that birds originated from bipedal long-tailed cursorial reptiles 
which during running oared along in the air by flapping thew 
free anterior extremities. It Dinosaurs had bird-like pulmonary 
appendages, as indicated by the pneumaticity of the skeleton, such 
movement would only have been of advantage for the respiratory 
organs (the pneumatic foramen occurring sometimes in Moa-bones 
would therefore be an atavistic feature, and the loss of pneumaticity 
would be a parallel to the same change in the Dinosaurian sub- 
class). At this point the pulmonary appendages of Chameleons 
have also to be taken into consideration. A double running and 
flapping action would—somewhat in accordance with Pycraft’s 
views on this subject—subsequently easily lead to an enlargement 
of the posterior marginal scales of the antibrachium, and at the 
same time produce a certain amount of bipedal specialisation. 

By gradually increasing in size, the enlarged but perhaps still 
horny hypothetical scales of the antibrachial margin would 

1907. ] ORIGIN OF FLIGHT. 235 

in time enable the yet carnivorous and cursorial ancestor of 
Birds to take long strides or leaps, much in the manner of a 
domesticated Goose or of a Stork when starting, and ultimately 
develop to actual feathers; this epidermic cover would also raise 
the temperature of the body, and thus help to increase the 
mental and bodily activity of these rapacious forms. The 
possibility of such a development of flight is clearly shown by 
the somewhat analogous, but still more marvellous and nearly 
paradoxical, yet not unfrequent, development of Flying- Fishes. 
The marginal scales being originally the principal wing-element 
in such a hypothetical form, these parts could attain quite a 
considerable size without essentially altering the underlying bones 
of the arm, a fusion of the carpal phalanges being only then 
necessary, when in flight rigidity of this region became requisite. 
Besides this, the continued use of the anterior flapping limbs as 

Text-fig. 82. 

Hypothetical reconstruction of a running “ Pro-Avis.” 

grasping-organs would also account for the feeble specialisation of 
the digits in the Ornitholestes-like manus of Archwopteryx, and 
for the preservation of the claws in the Ostrich and Opisthocomus, 
where, according to Pycraft, the temporary delay in the growth of 
the distal pinions has been developed simply not to prevent the 
claws from performing their still not unimportant function. An 
effort to condense these hypothetical changes into a drawing is 
given in text-fig. 82, which might in consequent allusion to 
Pyeraft’s analogous reconstruction be called a “ Pro-Avis.” 

The facts that even from the Hocene formation in most parts 
of the world numerous big Ratites are known, which can only 
have originated from badly-flying ground-birds, whereas in more 
modern times the Ratites are apparently vanishing from the 
earth’s surface, likewise find quite an easy explanation in the 
hypothesis that in the Mesozoic times badly-flying ground-birds, 
and not tree-birds, were the prevailing forms. The individual 
or ontogenetical development of every cursorial Carimate (for 
example, every gallinaceous bird) would thus show us the exact 
manner in which flight has been acquired. The true phylogenetic 
value of the surviving ‘‘ Paleeognathee,” with their body-temperature 


decidedly lower than in other birds on the one hand (Sutherland), 
and their reduced brachiosternal muscles on the other (Fiirbringer), 
can likewise be appreciated only if we consider them as forms 
that specialised at a very early stage of Avian evolution. 

It is to be remarked that among the terrestrial birds which 
according to this hypothesis would seem to have preserved their 
original mode of living and manner of breeding, the nest-building 
faculty is less developed than in those birds which, to avoid 
the dangers of ground-life, migrated up into the trees and had 
then to shelter their eggs and young ones from the new chance 
of falling to the ground. That ground-life involves for a bird 
more dangers than life on a tree, I think, is shown by the fact 
that the true ground-birds usually are protectively coloured, while 
in the latter, even among Gallinaceous Birds, bright—one might 
nearly say artistic—sexual characters are frequently developed. 

The supposition, that Birds once possessed a patagium and only 
afterwards developed feathers, I consider as devoid of foundation, 
for apart from the impossibility of a marginal feather being 
effective, when only attached to a flexible membrane, it is loss 
and not development of hair and scales (= epidermal coverings) 
that takes place in the Chiropterygian and Pterosaurian patagium. 
Besides, I do not see any reason why a useful patagium, once 
developed, should suddenly have stopped growing. 

The long tail in Archwopteryx can in no way be invoked in 
favour of a primitive arboreal stage of Birds, for a long tail not 
only characterises arboreal but also bipedal cursorial and saltatorial 
forms. Thus we cannot find a single character in Archeopteryx that 
would absolutely prove arboreal specialisation, while the develop- 
ment of the cannon-bone alone is sufficient to show with certainty 
that some of the direct ancestors of Archwopteryx had cursorial 


From a consideration of the whole of the above remarks, we 
can, I believe, formulate the following statement :— 

While Pterosaurs and Bats originated independently from 
quadrupedal arboreal forms in which both anterior and posterior 
extremities, in consequence of the development of a patagiwm, 
hecame primarily equally used for flight and in consequence equally 
unfit for locomotion on the ground, Birds originated from bipedal 
Dinosaur-like running forms in which the anterior extremities, on 
account of flapping movements, gradually turned to wings without 
thereby affecting terrestrial locomotion. This 2s also the reason why 

Birds became dominant over all the rest of their aerial rivals. 

In conclusion, I take pleasure in thanking once more all the 
ventlemen that helped me to compile this paper, notably Dr. K. 
Andersen, Mr. G. A. Boulenger, Dr. Forsyth Major, Mr. W. P. 
Pyeraft, and Dr. A. 8. Woodward, at the British Museum of 
Natural History. 

P.Z.S. 1906, pp. 759-1052 were published on April 11th, 1907. 

No. 38. 

January 15th, 1907. 

Dr. J. Rost Braprorp, F.R.S., Vice-President, in the Chair. 

The Secretary read a report on the additions that had been 
made to the Society’s Menagerie during the months of November 
and December 1906. 

Mr. Ouprretp THomas, F.R.S., exhibited the skin of a new 
Monkey from the Ituri Forest, obtained during the recent Ruwen- 
zori Expedition, and diagnosed it as follows :— 


Allied to C. campbell, but not darkened on the posterior back 
and hind limbs, the outer side of these latter being grizzled olive- 
yellowish to the ankles. Under surface and inner side of limbs 
very sharply defined creamy white. Tail greyish white, darkening 
terminally to black. 

Hab. Ituri Forest. 

Type. Male. Original number 184. Collected by R. E. Dent. 

Mr. P. H. Banr, F.Z.8., read a paper “On the ‘ Bleating’ or 
‘Drumming’ of the Snipe (Gallinago celestis).” The object of 
the paper was to prove that this phenomenon was produced by 
the tail-feathers of this species, a point which had been much 
disputed. It was found that if the feathers were attached to a 
cork in a special manner, the peculiar bleating sound could be 
produced, and, furthermore, that only two feathers in this species 
were the active agents in producing the sound. Observation 

* This Abstract is published by the Society at 3 Hanover Square, London, 
W., on the Tuesday following the date of Meeting to which it refers. It will 
be issued, free of extra charge, to all Fellows who subscribe to the Publications, 
along with the ‘ Proceedings’; but it may be obtained on the day of publication 
at the price of Sixpence, or, if desired, sent post-free for the sum of Six 
Shillings per annum, payable in advance. 


proved that these two feathers were held in a particular manner 
in front of the others during the bird’s flight in the breeding- 
season. Feathers of both male and female were found to bleat, a 
fact which had been borne out by numerous observers in the field. 
These feathers were found to have a peculiar structure, differing 
materially from the other feathers in the tail. Microscopically 
they differed, and the number of hamuli were found to be in excess 
of those found in other feathers. The feathers of various exotic 
species had been experimented upon, and those of G. delicata, 
nobilis, frenata, paraguaye in the New World, G. australis and 
aucklandica in the Antipodes, and G@. solitaria and megala in 
Asia had been found to produce musical sounds. These feathers 
varied in structure, and consequently the sound produced differed 
accordingly. The feathers of G. gallinula, G. major, and G. stenura 
were not found to be musical. 

Mr. J. L. Bonnote, F.Z.S., communicated a paper on a collec- 
tion of Mammals from Annam sent home by Dr. Vassal. Twenty- 
four species were enumerated, of which the following four were 
described as new :— 


Similar in general colouring to, but about half the size of, 
WV. coucang, and without any dark ‘markings round the eyes or 
down the back. The teeth are quite different, the second molar 
being the largest, whilst the third molar is triangular in shape 
and but little inferior in size to the first. 


Similar in general colouring to 7’. belangeri, but larger, with a 
much thicker tail and lacking the light neck-stripe. 


Similar to S. leucopus, but darker, and the outer sides of the 
limbs concolorous with the back. 


Similar to /. r. typicus, but much darker in general coloration 
and having a rufous tinge on the outer sides of the thighs. 

A paper was read from Dr. Emin A. Gorxp1, C.M.Z.S., contain- 
ing descriptions of seven new or little-known species of Marmoset 
Monkeys from the Amazonian Region. 

Mr. F. E. Bepparp, F.R.S., read a paper entitled “ Contribu- 
tions to the Knowledge of the Systematic Arrangement and 
Anatomy of certain Genera and Species of Squamata.” 

A communication was read from Mr. Groree H. Kenricsg, 
F.Z.S., containing a list, with descriptions of the new species, of 
Pyralide collected by Mr. A. E. Pratt in British New Guinea in 

The next Meeting of the Society for Scientific Business will 
be held on Tuesday, the 5th February, 1907, at half-past Hight 
o'clock p.m., when the following communications will be made :— 

1. Prof. E. Ray Lanxsster, F.R.S.—On the Feetus of the 

2. Dr. W. T. Caiman, F.Z.8.—On new or rare Cumacea from 
the Collection of the Copenhagen Museum. Part I. 

3. Dr. HE, A. Goetp1, C.M.Z.8.—Description of a new Amazonian 
Tree-Frog with peculiar Breeding-habits. 

The following Paper has been received :— 

Mr. C. J. Wirn.—An Account of the South-American Cheli- 
Jerine in the Collections of the British and Copenhagen Museums. 

Communications intended for the Scientific Meetings of the 
ZOOLOGICAL Society oF Lonpon should be addressed to 


3 Hanover Square, Lonpon, W. 
January 22, 1907. 

No. 39. 



February 5th, 1907. 

H.G. Tue DuxKE or Beprorp, K.G., President, in the Chair. 

Mr. F. Martin Duncan, by permission of the Charles Urban 
Trading Co., Ltd., gave a cinematograph exhibition of animals in 
the Society’s Gardens and other Zoological Studies, chiefly on the 
life-history of Insects. 

Mr. Ouprietp Tuomas, F.R.S., exhibited a collection of 
Mammals and Birds from the Islands of Saghalien and Hokkaido, 
N. Japan, made by Mr. Malcolm P. Anderson in carrying out the 
Duke of Bedford’s Exploration of Eastern Asia. Mr. Thomas 
proposed to give a full account of the Mammals on a later 

Mr. Oupriztp Tuomas also read a paper on Mammals collected 
in Mindanao, Philippines, by Mr. M. P. Anderson for the Duke 
of Bedford’s Exploration of Eastern Asia. Seven species were 
mentioned, of which the following was new :— 


Rather larger than C. fallax, of Luzon. Colour wholly dark 
blackish brown, not lighter below; limbs and tail also uniformly 

Dimensions of type:—Head and body 98 mm.; tail 68; hind 
foot 25. Upper molar series 4:1. 

Hab. Mt. Apo, Mindanao. Type. Male. No. 751. 

* This Abstract is published by the Society at 3 Hanover Square, London, 
W., on the Tuesday following the date of Meeting to which it refers. It will 
be issued, free of extra charge, to all Fellows who subscribe to the Publications, 
along with the ‘ Proceedings’ ; but it may be obtained on the day of publication 
at the price of Sixpence, or, if desired, sent post-free for the sum of Sic 
Shillings per annum, payable in advance. 


Prof. KE. Ray Lanxester, F.R.S., Director of the British Mu- 
seum (Natural History), read a paper entitled “The Origin of the 
Lateral Horns of the Giraffe in Foetal Life on the Area of the 
Parietal Bones.” The author described and showed the exact 
relation of the lateral horns in the fetus taken from the Giraffe 
which died last spring in the Society’s Gardens. It was demon- 
strated that the lateral horn of the Giraffe was exclusively in 
origin a part of the fibrous osteogenetic tissue of the parietal bone 
of which it was a part, and had no connection whatever with the 

Thus the statement made by Su: Richard Owen in his account 
of a new-born Giraffe, in a paper read before the Society in 1839, 
was finally shown to be based on an unfortunate accident. Owen 
had cut out the horn-bearing area of the skull and after an interval 
of time had reversed the relations of the excised piece of bone, 
taking frontal for parietal and parietal for frontal. 

The author expressed the opinion that the parietal lateral horn 
of the Giraffe could not be considered to be the same morphological 
unit as the frontal lateral horn of the Okapi. 

Prof. LANKESTER also read a paper on “ Parallel Hair-fringes 
and Colour-striping on the Face of Foetal and Adult Giraffes,” in 
which he described, illustrated by lantern-slides, a remarkable 
eolour-banding or striping of the hairy covering of the face in the 
foetal Giraffe, and showed that similar dark and light striping 
occurred in a very marked form in adult Giraffes though not in 
all individuals. 

Ina third paper, ‘On the Existence of Rudimentary Antlers in 
the Okapi,” Prof. Lanxester described the polished tip or apex of 
the Okapi’s horn which breaks through the integument. He 
showed that transverse fissures or incisions were produced one 
behind the other in the naked apex, tending to cut off in succession 
a series of small bony caps which he regarded as rudimentary 
antlers. He expressly refrained from concluding that this forma- 
tion of minute antler-caps was to be regarded as genetically 
connected with the antler-formation of the Cervide, though such 
a connection was possible. 

Prof. LANKESTER also exhibited the skull of a sub-adult male 
Okapi obtained by Major Powell-Cotton in the Ituri Forest 
(Congo), and a similar somewhat younger male skull, obtained by 
Capt. Boyd Alexander in the Bahr-el-Ghazal region. Both skulls 
were in a very fine state of preservation. 

Mr. G. A. Bountencer, F.R.S8., on behalf of Dr. E. A. GorELp1, 
G.M.Z.S., exhibited and described a new Amazonian Tree-Frog, 
Hyla resinifictrix, closely related to H. venulosa, but distinguished 
by fully half-webbed fingers. This frog was remarkable for its 
habit of making good-sized basins of resinous substances in hollow 
branches of high trees, in which water collects, which served as a 


nursery for the eggs and larve. The fzog collected the resin 
from the bark of certain trees, such as the aromatic “brew- 
branco ” (Protiwm heptaphyllum). 

Dr. W. T. Catan, F.Z.S., read the first part of a paper on the 
Collection of Cumacea in the Copenhagen Museum. Altogether 
30 species were dealt with, of which 25 were described as new. 
The majority of the specimens were derived from collections made 
in New Zealand and the Gulf of Siam by Mr. H. Suter and 
Dr. Th. Mortensen respectively. 

The next Meeting of the Society for Scientific Business will 
be held on Tuesday, the 19th February, 1907, at half-past Hight 
o'clock P.M., when the following communications will be made :— 

1. Mr. R. I. Pocock, F.Z.8.—On English Domestic Cats. 

2. Dr. C. G. Seniemann, F.Z.S.—On Deaths occurring in the 
Society's Gardens during 1906. 

3. Mr. J. T. Cunnincuam, F.Z.5.—On a peculiarly Abnormal 
Specimen of the Turbot. 

4, Baron F. Nopcsa.—Ideas on the Origin of Flight. 

The following Papers have been received :— 

1. Mr. C. J. Wrrn.—An Account of the South American Cheli- 
Jerine in the Collections of the British and Copenhagen Museums 

2. Dr. R. Broom, C.M.Z.S.—On the Dental Succession in the 
Cape Golden Moles, 

3. Mr. F. E. Bepparp, F.R.S.—On the Azygos Veins in the 

4, Mr. F. E. Bepparp, F.R.S.—On Two new Species of the 
African Genus Microchetus belonging to the Collection of Oligo- 
cheta in the Museum of Christiania. 

5. Miss Dorotuea M. A. Bate.—On Elephant Remains from 
Crete, with Description of Hlephas ereticus, sp. nov. 

6. Mr. Cuaruzs F. Roussever.—Zoological Results of the Third 
Tanganyika Expedition, conducted by Dr. W. A. Cunnington, 
1904-05. Report on the Polyzoa. 

Communications intended for the Scientific Meetings of the 
ZOOLOGICAL SocIETY oF Lonpon should be addressed to 


February 12, 1907. 

No. 40. 



February 19th, 1907. 

Sir Kpmunp G, Lopzr, Bt., Vice-President, in the Chair. 

The Secrerary read a report on the additions that had been 
made to the Society’s Menagerie during the month of January 

Dr. C. I. Forsyra Magsor, F.Z.S., exhibited remains of a Bear 
from the superficial deposits of a cavern in the mountains of 
Corsica, where Bears, though now extinct, were formerly nume- 
rous, at least up to the sixteenth century. Despite the fact that 
no truly fossil Bears were as yet known from Corsica, Dr. Forsyth 
Major considered the Corsican Bear to have been autochthonous, 
whilst in his opinion the recent Mammals of Corsica (and Sardinia) 
had been, almost without exception, introduced by human agency. 
In any case they could not be adduced as proofs of a recent con- 
nection of those islands with either of the neighbouring continents. 

In a paper on English Domestic Cats, Mr. R. I. Pocock 
urged that the surest basis for their classification and the most 
satisfactory clue to their descent was furnished by the two dis- 
tinct patterns found in so-called Tabby Cats. In one type the 
pattern consisted of narrow vertical stripes ; in the other of longi- 
tudinal or obliquely longitudinal stripes which, on the sides of 
the body, tended to assume a spiral or subcircular arrangement 
characteristic of the “blotched” Tabby. This distinction was 
long ago pointed out by Blyth. 

One or the other of these types was to be found in Cats of almost 
all breeds, whether “ Persian,” ‘‘ Short-haired,” or “ Manx.” There 
appeared to be no intermediate stages between the two. The Cats 
of the “striped” type were no doubt descended from the Kuropean 
Wild Cat and the North-African Wild Cat; but the origin of 

* This Abstract is published by the Society at 3 Hanover Square, London, 
W., on the Tuesday following the date of Meeting to which it refers. It will 
be issued, free of extra charge, to all Fellows who subscribe to the Publications, 
along with the ‘ Proceedings’ ; but it may be obtained on the day of publication 
at the price of Sixpence, or, if desired, sent post-free for the sum of Six 
Shillings per annum, payable in advance. 


Cats exhibiting the “ blotched” pattern appeared to be unknown. 
It was to the Cat of the latter kind that Linneus gave the name 
catus, which was therefore no longer available for the European 
Wild Cat; this Cat, therefore, must take the name sylvestris. 

Dr. C. G. SznieMANN, the Society’s Pathologist, in presenting 
his report on the deaths that had occurred among the Mammals 
and Birds in the Menagerie during 1906 stated that 356 Mammals 
and 283 Birds were submitted to post-mortem examination, and 
the results showed— 

(i) That tuberculosis occurring in birds in the Gardens was 
usually due to infection by the gut. 

(ii) The hearts of Rheas, Cassowaries, Ostriches, and some 
of the larger Storks kept in the Gardens were often 
extremely flabby, and death in these birds was in a 
large number of cases due to cardiac failure. 

(iii) New growths were rare both in mammals and in birds, 
but one case of carcinoma arising in the kidney and 
occurring in a Chilian Pintail (Dajila spinicauda) had 
been observed, as well as two instances of benign new 
growths occurring in birds not inmates of the Gardens. 

Mr. J. T. Cunninenam, M.A., F.Z.8., described a peculiarly 
abnormal specimen of the Turbot. The specimen was captured 
by Miss Olivia Fox, of Falmouth, near Padstow, on the north 
coast of Cornwall. It was a young fish, measuring only 4°4 cm. 
in length, and a normal specimen of slightly smaller size, taken at 
the same time, was completely metamorphosed to the asymmetrical 
condition of the adult. In the abnormal specimen the right side 
was almost entirely destitute of colour as in the normal condition, 
but both eyes were on this white side, instead of being on the left 
side as in normal Turbot. On the left side pigment was present 
over the whole surface except the head and the anterior part of 
the base of the dorsal fin, which were white. The fish was kept 
alive in captivity for two months, and was observed to lie always 
with its eyes uppermost, so that the upper side was white and the 
lower side coloured. The fish showed also another abnormality, 
namely, that the base of the dorsal fin projected anteriorly as a 
free process above the dorsal eye, a peculiarity which is usually 
present in ambicolorate Turbot. As there was some pigment on 
the head on the left side, Mr. Cunningham pointed out that the 
specimen might be regarded as a Turbot in which a normal body 
was united with a head which was reversed, so that the left side 
of the head, bearing the eyes and pigment, was joined to the right 
side of the body bearing no pigment, and vice versa. 

Dr. Baron Francis Norcsa read a communication entitled 
“Ideas on the Origin of Flight,” and illustrated his argument 
with lantern-slides showing the hind limbs of various genera of 
Bats, Pterosaurs, Birds, and Dinosaurs, as well as a reconstruc- 
tion of a hypothetical, cursorial primitive bird. The author 


stated that from the mechanical point of view a patagium and a 
set of flight-feathers were different organs. He pointed out the 
osteological analogies between Bats and Pterosaurs on the one 
hand, and between Birds and Dinosaurs on the other. He sug- 
gested that Bats and Pterosaurs had arisen from leaping, arboreal 
forms, whilst Birds had come from a terrestrial, cursorial stock. 

Mr. F. KE. Bepparp, F.R.S., read a paper on the Azygos Veins 
in the Mammalia. 

The next Meeting of the Society for Scientific Business will 
be held on Tuesday, the 5th March, 1907, at half-past Hight 
o'clock P.m., when the following communications will be made :— 

1. The Hon. Watter Roruscuinp, M.P., F.Z.8.—Descriptions 
of some new Species and Subspecies of Antelopes and of a new 

2. Miss Dorornea M. A. Barr.—On Elephant Remains from 
Crete, with Description of a new Species. 

3. Mr. Cuaruses F, Roussever.—Zoological Results of the Third 
Tanganyika Expedition, conducted by Dr. W. A. Cunnington, 
1904-05. Report on the Polyzoa. 

4, Dr. W. A. Cunnineton.—Zoological Results of the Third 
Tanganyika Expedition, conducted by Dr. W. A. Cunnington, 
1894-05. Report on the Brachyurous Crustacea. 

The following Papers have been received :— 

1. Mr. C.J. Wirn.—An Account of the South American Cheli- 
ferine in the Collections of the British and Copenhagen Museums. 

2. Dr. R. Broom, C.M.Z.S.—On the Dental Succession in the 
Cape Golden Moles. 

3. Mr. F. E. Bepparp, F.R.S.—On Two new Species of the 
African Genus Microchetus belonging to the Collection of Oligo- 
cheeta in the Museum of Christiania. 

4, Mr. T. A. Cowarp, F.Z.S.-—On the Winter Habits of the 
Greater Horseshoe and other Cave-haunting Bats. 

Communications intended for the Scientific Meetings of the 
ZooLoGicaL Society or Lonpon should be addressed to 


3 HANovER Square, Lonpon, W. 
February 26, 1907. 

is 5) * Fy ees i : ys RRS) Pen gg ase 
pene Uy ooaaee Lata gine i 

awe. i 


) nM 

tt ie eft ; ‘wh 

be eee bY) 


oi, Hid Puilenuetea 



" fale eae 

+h aren 

Contents (continued). 

February 5, 1907 (continued). 

Va ' : Page 

Dr. W. T. Calman, F.Z.S. Notice of a paper on New or Rare Crustacea of the Order 
Cumacea from the Collection of the Copenhagen Museum ....-.......+204-.ss000% 100 

1. The Origin of the Lateral Horns of the Giraffe in Foetal Life on the Area of the Parietal 

Bones. By E. Ray Lanxesrer, M.A., D.Sc., LL.D., F.R.S., F.Z.8., Director of the 
Natural History Departments of the British Museum ........002,..--00-+--0e-- 100 

2. Parallel Hair-fringes and Colour-striping on the Face of Fetal and Adult Giraffes. 

By E. Ray Lannester, M.A., D.Sc., LL.D., F.R.S., F.Z.8., Director of the British 
Mobseun (Natural History): (Plate V.) 2. ecco. str el wierspets & nbalebaye) eiehalg) a elthele Vere ete 115 

3. On the Existence of Rudimentary Antlers in the Okapi. By E. Ray Lanxssrer, M.A., 

D.Sc., LL.D., F.RS., F.Z.8., Director of the British Museum (Natural History). 
Gialeipes VB Gc VEL NS ge gees bas 5.0 4 siath does siscaig oco'els hc 2S sie ay rep er 126° 

4. Description of Hyla resinifictriz Goeldi, a new Amazonian Tree-Frog peculiar for its 

Breeding-habits. By Prof. Dr. Emm A. Gorupr, C.M.Z.S., Director of the Para 
AVEUT SESE S2o:\cccvs' vie si aiel al ip'ai ='n wa a\sialsfalaiereisin<cisvale/e\sye & « lele:o/m\egeleiphiie: tet uhe a/carete <laaaeaia mens 135 

5, The Duke of Bedford’s Zoological Exploration in Eastern Asia.—lIT. On Mammals 
obtained by Mr. M. P. Anderson in the Philippine Islands. By O:prrety Tuomas, 

PEE Bose ar ols lc) Salelg initials a sip nia ci ejoie w\nisie\n's e's wiv s d.suslelnh ie ula shat el oiler ke tees 140 
February 19, 1907. - 
The Secretary. Report on the Additions to the Society's Menagerie during the Month of 
Uae US, CEBS 6.46 acto Ap POCO EO BEIEEGBRDe ORDO S a6 go bininieUie'a sie’e) jain cial ama ae etage 143 
Dr. C. I. Forsyth Major, F.Z.S8. Exhibition of remains of a Bear from a Cavern in 
WOPSIEAS azetaiec « < foe) Sees Sot EGA ee CDR Ones a oabtds. Suopeouesaoéancass . - 143 
1. On English Domestic Cats. By R. I. Pococx, F.L.S., F.Z.8., Superintendent of the 
Zoological Society’s Gardens. (Plates VIII.-X.) ..-............ picket iwietasyorenyaiats 145 
2. Report on Deaths occurring in the Society’s Menagerie during 1906. By C. G. Sztiamayn, 
M.D, ) We See atholorist to-the Society... cteic <n: sce ve «0 secs elase areas catahsleeer aaa 168 
3. On a peculiarly Abnormal Specimen of Turbot. By J. T. Cunninenam, M.A, F.Z.S. 
(CREW: GI ERC APC a ie Ce NIE A ERDAS Coinenmitiy dik GOO HME OO AR eke ac 174 
4, On the Azygos Veins in the Mammalia. By Frank E. Bepparp, M.A. (Oxon.), 
F.R.S., Prosector to the Society ........... Rcletsietaratatel ets duct acarevefats Map uO Bente eeitc a Lenk 

. Ideas on the Origin of Flight, By Dr. Baron Fraxcis Norcsa ..- . 



1907, pp. 1-236. 

Plate ; Page 
BES COncOptBECUS DOMUE Svan e's au So's sao baie: csis Sayers enh 2 
TD VEVELCCOD US UOINUS e's sia «0's, bicie's ce ein mnlensl eee ene ele 3 
IV } Pyralidx from British New Guinea ...............-00 - 68 
V. Head of a Fetal Giraffe (half the natural size) ........-.. 115 
VI. Horn-tips of Okapi ..... BAA Aan aaah dole AON ItoS b's 4 a 
VII. Young Ossicone or Horn of Okapi .,..........-++.00..-- } eas 
VILE.” Blotched Tabby, Felis catus 2... sie ce vee oo eens ee | 
PX Striped: Pabby, Hales Conguata i. « cues cose ok mica eeie iets 143 
X. Agriotypes of the Striped Tabby ...... Bea) Sead aah J 
xt.) Abnormal Younr: Parbobs. 6s tah 5 oss os = gi + chet lseiecle aii me 

The ‘ Proceedings’ for the year are issued in four parts, forming two volumes. 
as follows:— 
Papers read in January and February, in June. 
i » March and April, in August. 

May and June, in October. 

” ” 

x » November and December, in April. 

‘ Proceedings,’ 1906, pp. 759-1052, were published on April 11th, 1907. 

The Abstracts of the papers read at the Scientific Meetings in 
January and February are contained in this Part. 







Paces 237-446. 


AUGUST 1907. 



| Yas [Price Twelve Shillings. ] 
bees é ( 

March 5, 1907. 

The Hon. Walter Rothschild, M.P., F.Z.S. Exhibition of a mounted specimen of a 
Grama e re ee is a eek ots Ge ms spe rahe age By aed Ded ie HEI AN deednat D's oll 237, 
1. Descriptions of a new Species‘and two new Subspecies of Antelopes and a new Sheep. 
By the Hon. Waurnr Roruscaiyp, M.P., Ph.D., PREZ Gere ctene, Cie iSl siete enere ante cneent ene £237 
2. On Elephant Remains from Crete, with Description of Hlephas creticus, sp.n. By 
Dorornma M. A. Barn. (Plates XIT. & XTIL.) .. ee eee eee eee ees witha Ale game 
,9. Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A. 
Cunnington, 1904-1905.—Report on the Polyzoa, By Cuaruns FE. Rovussexer, 
EVR.M.S. . (Plates XTV. & KV.) cece cece ee cere eee cet ees Bie see astanan aa Soilt cet 250° 
4. Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A 
Cunnington, 1904-1905.—Report on the Brachyurous Crustacea. By Winwiam A.” 
Cunninoron, B.A., Ph.D., F.Z.S. (Plates XVI. & XVIL.)...- ee eee ee eee ee eee 208 
5. On Two new Species of the African Genus Microchetus belonging to the Collection of , 
Oligocheta in the Museum of Christiania. By Frank E, Bepparp, M.A., F.RS,, 
Prosector to the Society ........ esse ee ee ee ee ee ee es BA IOS Us NE a eee Ea QT 

March 19, 1907. 

The Secretary. Report on the Additions to the Society's Menagerie during the month of 
February 1907.2... ce cece cee ee tener eens Bea chat obs calanecalcale fe] cuss catetote 281 

-Mr. Herbert F. Standing. Notice of ‘a Memoir on recently discovered Subfoussil Prosimiz 
from Madagascar ........ SEN APT ER NADL OR EAL gisem cunts AAG et --- 28h 

1. Descriptions of some New Species of Animal Parasites. By L. W. Samson, M.D., HAS. 282 

2. Descriptions of five New Species of Hamogregarines from Snakes. By L. W. Sampon, 
M.D., F.Z.S., and 0, G. Seniamann, M.D., F.Z.8. ........ 0 Ph Reape ec Sees 28 

3. The Rudd Explor ation of South Afvica,—VII. List of Mammals obtained by Mr. Grant 
at Coguno, Inhambane. By Oxpriztp Tuomas, F.R.S., F.Z, S,,and R. C. Wrovenron, 
1S AIS Ce er a @eneese ae beapeeeeereenere ee een enews ee ey 285 

Contents pomied on page 3 of Wrapper. 


March 5, 1907. 
Freperick Giniert, Ksy., Vice-President, in the Chair. 

The Hon. Walter Rothschild, M.P., F.Z.S., exhibited a mounted 
specimen of a Gorilla, Gorilla gorilla diehli. 

The following papers were read :— 

1. Descriptions of a new Species and two new Subspecies of 
Antelopes and a new Sheep. By the Hon. Waurer 
RorascHiLp, M.P., Ph.D., F.Z:8. 

[Received March 3, 1907.} 


Differs from 2. horstocki in its much darker colour. The white 
patches on the throat and round the eyes much smalier and the 
white on the belly less extended. The whole of the rest of body, 
head, and limbs dark vinaceous rufous instead of orange-rufous. 

Hab. Neighbourhood of Drakensberg, Natal. 


Differs from C. arundinewm in the paler, more greyish rufous 
of head and neck and the pale rusty-grey of limbs, tail, and body. 
The horns also seem straighter and thicker as a rule. 

Hab. Neighbourhood of Fort Jameson, Northern Rhodesia, and 

Bangweolo Flats. 

CoBuUs ROBERTSI, sp. 0. 

IT am treating this as a species because it occurs in the same 
territory as true C. lechwe and C. smithemanni, otherwise it has 
many characters intermediate between these two species. Nor 
do I consider it likely that it is a hybrid, for it has been shot by 
several sportsmen, and not always in the same localities. 

Horns stouter and the rings broader and closer together than 
in ©. lechwe, smooth tip shorter. Head, hind-neck, body, and 
upper part of fore-neck bright orange-rufous, back and flanks 
darker and redder than in C. lechwe. Cheeks, throat, and sides 
of neck have the rufous strongly mixed with black hairs, thus 
somewhat approaching C. smithemanmi. Lower fore-neck whitish, 
with two large black patches on each side, which join similar 
patches on the front of the shoulders. Outside of ears rafous 

buff, inner side white. 
Hab. Northern Rhodesia. (Type, collected by Mr. Roberts.) 

Proc. Zoou. Soc.—1907, No. XVII. AF 

238 MISS D. M. A. BATE ON [ Mar. 5, 

OvIs COWANI, sp. n. 

This Sheep is nearest to O. stonrei, but differs in being entirely 
deep black, with the exception of a white rump-patch and a grey 
face. The rump-patch is considerably smaller than in O. stonet. 
The three-year old ram of O. stonei is occasionally much darker 
than a specimen of that species in my possession, but is always 
of a rusty or brownish black, and has the very large white rump- 

The type specimen of O. cowani was entire, but the skin had 
only been dried and not dressed, and when relaxed the whole of 
the hair came off, and only the head and neck could be saved. 

The type, shot by an Indian out of a large flock, was sent 
me by Mr. C. G. Cowan, of Kamloops, and was obtained in the 
mountain-chain near Mount Logan in British Columbia. 

Types of all these four species and subspecies are in the Tring 

2. On Elephant Remains from Crete, with Description of 
Elephas creticus, sp.n. By Dorornea M. A. Bats.* 
[Received February 1, 1907. | 
(Plates XII. & XIII.7, and.Text-figure 83.) 


Perhaps the most important and interesting of the results of 
the author’s visit. to Crete in 1904 was the discovery, in two 
hitherto unexplored cave-deposits, of the remains of Elephants of 
different sizes, more particularly as the occurrence of one of these, 
of pigmy proportions, appears not to have been previously known. 

Although prior to 1904 no large quantity of specimens seems 
to have been obtained, yet the existence of ossiferous deposits 
in this island has been known for a considerable time, as the 
following records testify. The earliest would seem to be that of 
Pococke, who described a bone-cave in the Khania Akrotiri in a 
volume published in the middle of the eighteenth century 2. 
Nearly a hundred years after this, a reference occurs relating to 
fossilised human remains found, together with marine forms, near 
Khania by Fabrequette §, who was at one time Consul at Malta. 
Later, remains of a Hippopotamus, which prokably came from 
the upland basin of Lassethe, were obtained by more than one 
traveller ||, and have since been referred to by a number of writers. 
Two ossiferous caves were discovered in the west of the island by 

* Communicated by Henry Woopwarp, LL.D., F.R.S., V.P.Z.S., F.GS. 

+ For explanation of the Plates, see p. 250. 

+ Richard Pococke, ‘A Description of the East’ (London, 1745), vol. 11. p. 264. 

§ C. R. Acad. Sci. (Paris), iv. 1837, p. 182; also ibid. viii. 1839, p. 178. 

|| See Admiral Spratt. ‘Travels aud Researches im Crete’ (London, 1865), vol. 11. 

pp- 386-7; also Raulin, ‘ Description Physique de l’Ile de Créte’ (Paris, 1869), vol. i. 
p. 156, and vol. 11. p. 615. 

Das, WOW, IPL, 20. 



West,Newman imp. 

G.M-Woodward lith. 




-durr weurms yy “4Sse “UQIT 99" TOpP Puempoom WD 

“TINDSC Weal, AA OWS SIE TS) 74 val 


Admiral Spratt *, both of which have since been visited by the 
present writer ‘. 

Signor Simonelli, whose geological researches in Crete were 
carried on in 1893, was seemingly the first to obtain Elephant 
remains, which he procured from cayes near Retymno, which is 
situated on the north coast and les between Khania and Candia. 
In a paper published in 1894 = he identifies the specimens, which 
included a perfect mandibular ramus, as those of Elephas priscus 
(=4. antiquus §), though no detailed descriptions or figures are 
given, and I have been able to find no other published reference 
to them. 

Unfortunately the collection of teeth and limb-bones which 
forms the subject of the present paper consists of a small amount 
of material only, many of the specimens being but imperfectly 
preserved. Notwithstanding this they fall naturally into two 
groups representing animals of different sizes, the smaller of 
which agrees in this respect with H. melitensis, whilst a number 
of limb-bones indicate an Elephant superior in dimensions to the 
average Indian species (#7. maximus) of the present day, but not 
equalling the gigantic proportions at times attained by Z. meri- 
dionalis and L. antiquus, though, as will be shown later, it is to 
this last that they must evidently be referred. Thus at first sight 
it seemed probable that the collection included remains of both a 
dwarted race and the parent form from which it had sprung. 
However, before remarking on the relationships suggested by a 
study of the specimens obtained, it will perhaps be more con- 
venient first to briefly describe them, commencing with those by 
which the pigmy form is represented. 


The remains of the smallest of the Cretan Elephants were all 
obtained from a much damaged and weathered cave-deposit in the 
limestone cliffs near Cape Maleka in the west of the island, which 
has already been described ||, and where only some teeth and 
limb-bones of small rodents were found besides those under dis- 
cussion. These latter include nine imperfect molars and a few 
fragments, among which are a portion of an incisor and the 
dorsal half of a vertebra. As this small race differs from those 
of other Mediterranean islands, and its minute proportions being 
seemingly the result of specialisation due to isolation in Crete, it 
is suggested that it may be known by the above specific name 
denoting its island habitat. 

Incisors—Of the milk-incisors no specimen was _ procured, 
while, as already remarked, only a fragment of the proximal end 
of a permanent tusk (M. 9375) was obtained. From the limited 

* Op. cit. vol. u. pp. 194-5. 

+ Geol. Mag. n. s. dec. v. vol. 11. (1905) pp. 194-6. 
t Atti R. Accad. Lincei, 5* ser. Rendiconti, vol. i. 1894, Sem. 2, pp. 265, 268 ; 

also ‘Candia’ (Parma, 1897), pp. 171-2. 
§ Cat. Foss. Mamm. Brit. Mus. part iv. (London, 1886) p. 122. 

|| Geol. Mag. n. s. dee. v. vol. 11, (1905) p. 195. 

240 MISS D. M. A. BATE ON [ Mar. 5 

extent of the pulp-cavity, it is believed to have belonged to a nearly 
adult individual and indicates a tooth of small size, though its 
condition is too imperfect to admit of any measurements being 
given other than its approximate diameter, which is 46 mm. 

Upper Molars. ——(Ornlhy two of the nine sanvellanis are considered to 
belong to the upper series; the smaller of these (M. 9376), a first 
true molar, i is split longitudinally aimost in half, but sufficient 
remains to show the general character of the tooth, It consists 
of seven plates, though it is possible that an additional small 
anterior ridge was originally Breseut: Its antero-posterior length 
is 54 mm, and height about 32 mm., although the fangs s extended 
for a short distance further, bringing the total measurement up 
to 41 mm, Comparatively speaking, the height of this tooth is 
slightly greater than in most of the other specimens referred to 
E. creticus; this is an argument in favour of its being a first true 
molar rather than the last of the milk- series, which, from its size 
alone, it might perhaps have been thought to represent. The 
enamel bands are not very thick and im several of the ridges are 
somewhat broken up into “rings”; at the same time, the cement- 
areas are broad, a character generally found in conjunction with 
more massive enamel. In section this tooth shows that the cement- 
areas maintain an almost uniform width for nearly the whole of 
their extent. This feature is probably not infrequent in the 
molars of this species, as suggestively exemplified by some of the 
extensively worn specimens described below. 

The Cretan specimen is superior in antero-posterior length to an 
example of the corresponding tooth of H. cypriotes (M. 8602) *, 
though this latter exceeds it in the height of its crown, which 
is 53-5 mm.; this difference in the proportions is even more 
marked in a molar of 1, melitensis (44252) r. 

The second specimen (M. 9577, PI. XCUL, fig. 2), believed to 
belong to the upper series, is of larger propor tions than the last, 
and is much worn and somewhat damaged ; it probably represents 
the penultimate true moiar. tis difficult to say of how many plates 
it originally consisted, but it can be seen that there were at least 
eight, all of which were in use, the front ones bemg worn down 
almost to their common base, while a portion of the anterior ridge 
is broken off. Partly owing to its advanced state of wear, the 
crown is very wide, being 37 mm. across, while its antero-posterior 
length is 77 min. and its height 37 mm., though this would 
of course have been greater when the tooth was still but slightly 
abraded. In spite of the condition of this molar the cement-areas 
are still broad, equalling or even exceeding in size the plates of 
dentine; the enamel bands are thick and in a few instances 
somewhat wavy in outline. 

Aithough apparently considerably superior in height of crown, 
this tooth in 4. cypriotes is otherwise decidedly smaller than the 
above, the dimensions of a much less worn example (M. 8601) 

* Fioured, Phil. Trans. vol. 197 B (1904), p. 351, pl. 21. figs. 4, 4a. 
+ Trans. Zool. Soc. vol. ix. p. 20, pl. 2. figs. 9, 9a. 

being :—approximate antero-posterior length 72 mm., greatest 
width of crown 28 mm., and total height 65 mm. Compared 
with the specimen (49267) * identified by Falconer as M. 1 of 
LE. melitensis, this last is somewhat narrower in the crown, which 
is 53 mm. across, though this may be partly accounted for by the 
difference in age, and it is likely that the two would originally 
have closely agreed in this respect. On the other hand, in height 
the molar from Malta greatly exceeds the one from Crete, being 
very high in relation to its width, as already noted by Falconer ; 
further, the cement-areas in the Cretan fossil are distinctly 

Lower Molars.-—Ot the seven molars obtained not one can be 
referred to a position among the milk-teeth, while, owing to 
imperfect preservation, it is impossible in one or two cases to make 
certain of the place originally occupied in the permanent series. 
The first true molar appears not to be represented. Two specimens, 
both belonging to the right side of the mandible, are believed 
to be second true molars. One (M. 9578 a) has been considerably 
weathered and is much worn, all except the last of the plates 
having been in use; the number of these was at least nine. The 
‘cement-areas are broad and the enamel bands thick and smooth, 
while in two, if not three, of them there is a median loop, 
though this is unaccompanied by a mesial expansion of the plates 
of dentine. 

The other tooth (M. 9378, Pl. XIIT. figs. 1, 1 a), regarded as a 
second true molar, is also extensively worn, the penultimate ridge 
being already slightly abraded ; although it is broken off anteriorly 
seven plates remain as well as a portion of an eighth. The width 
of the seventh plate (counting from the rear) is 37 mm., while the 
height of the preantepenultimate one is only 33 mm. : the enamel 
is massive and in the four posterior used plates is somewhat 
irregular, while the cement separating the anterior ridges is con- 
siderably excavated. A comparison shows this Cretan specimen 
to be broader and much lower in the crown than a corresponding, 
though less worn, tooth of £. eypriotes (M. 8588) 7 

Judging from their size it is probable that two other lower true 
molars, very imperfectly preserved, also represent the second of 
the permanent series. One of these (M. 9382) is situated in the 
posterior half of a left mandibular ramus; it probably consisted 
of eight or nine plates and, like most of the teeth obtained of 
E. creticus, is low-crowned and has thick enamel bands and wide 
cement-areas. The other example (M. 9380) is a portion of a much 
damaged right molar; the worn surfaces of two ridges remain and 
seem in no way to differ from those of the specimens already 

The last lower molar (M. 3) is represented by three specimens, 
none of which is in an advanced stage of wear. Two of these, 

* Falconer, Pal. Mem. vol. ii. pl. 11. figs. 2, 2a, p. 294; also identified by Busk as 
M. 1? of #. falconeri, Trans. Zool. Soc. vol. vi. part v. pl. 53. figs. 9, 9 a. 
+ Phil. Trans. vol. 197 B (1904), p. 355, pl. 21. figs. 3, 3a. 

242 MISS D. M. A. BATE ON [Mar. 5, 

damaged anteriorly, are each situated in a portion of a ramus; 
and belonging, as they do, to opposite sides of the mandible as well 
as being in a similar stage of wear, it is thought that they may 
have been owned by a single individual. On the left side 
(M. 9383, Pl. XII. fig. 1), in addition to the last molar, the 
three posterior plates of M. 2 are also present. In M. 3 eleven 
plates can be counted, but a considerable part of the posterior 
portion of the tooth is wanting. The fifth plate is slightly abraded, 
and it is noticeable in both this and the companion specimen that 
the enveloping cement on the grinding-surface of the crown is 
somewhat scanty, with the result that all the enamel soon becomes. 
exposed, even while the anterior portion of the tooth may still be 
but little worn. The second example (M. 9383 6) presents very 
similar characteristics ; the presence of eleven plates can be deter- 
mined, but damage anteriorly and concealment of the tooth behind 
py the boneand matrix make it impossible to ascertain the original 

The one other last true molar (M. 9381, Pl. XII. fig. 3), an 
isolated tooth of the left side, is almost perfectly preserved except 
for the loss of the greater part of the first and a portion of the 
second plate. It is considerably curved, this being accentuated by 
the angle at which the hinder plates lie. The number of ridges is 
thirteen, six of which show signs of more or less wear, and there 
appears to be no evidence of the former presence of any additional 
ones. The sloping, instead of upright, position of the last plate 
is a further indication that this molar must be considered an 
example of the last of the permanent series. The antero-posterior 
length of the tooth (not along the curve) is about 122 mm., or 
139 mm. if continued to the heel of its base; the greatest height, 
which occurs at the sixth plate, is 53 mm., while the width of the 
abraded surface of the third ridge is 3) mm. The cement-areas 
are of medium width and the enamel bands thick and uncrimped, 
though somewhat irregular and disconnected in the less-worn 

It is interesting to contrast this with two corresponding teeth 
of #. cypriotes in the British Museum Collection *, ail three being 
of the left side and in practically a similar stage of use. The 
Cretan fossil is the more massive and superior in width and 
antero-posterior length, the persistence of the annulation of the 
enamel bands is also more strongly marked. On the other hand, 
it has a lower crown (53 mm.), both actually and as compared 
with its bulk, than the Cypriote specimens, this being 59 mm. 
in. the isolated tooth (M. 8591), while in the other (M. 8589), 
although the ramus prevents a measurement of the greatest 
height being obtained, that of the seventh plate is 63 mm. 
The most satisfactorily identified last molar ef H. melitensis in the 
British Museum Collection for comparison with the above-noticed 
tooth from Crete, is a specimen situated in a right mandibular 

* M. 8589, Phil. Trans. vol. 197 B (1904), p. 355, text-figs. 2&3; and M. 8591, 
ibid. p. 355, pl. 22. figs. 6, 6a. 


ramus (M. 44294), already described and figured by Dr. Leith 
Adams*. It is evident that in antero-posterior length it 
slightly exceeds the Cretan example and consists of several more 
plates, these being in much closer proximity owing to the narrow- 
ness of the cement-areas and the less massive enamel bands. 
Considering the size of the ramus this tooth must be much the 
higher of the two, while in breadth of crown the molar from Crete 
is superior, 

The one other dental specimen obtained (M. 9379, Pl. XII. 
fig. 2) consists of only three unworn plates; the anterior surface 
of the foremost of these is strongly but simply grooved, its greatest 
width is 35 mm. and its height 50 mm. 

The only bone procured from the deposit near Cape Maleka 1s 
the dorsal half of a vertebra embedded in the matrix attached to 
the mandibular ramus containing a last true molar (M. 9383). 

From this brief account of the remains procured of H. creticus, 
it will be seen that this pigmy Elephant must have been of slightly 
larger proportions than #. cypriotes and approached in size more 
closely to H. melitensis ; that is to say, it would have attained as 
a maximum a height of five feet f. All the molars obtained differ 
from those of the two last-named dwarf species in being much 
lower in the crown; this is perhaps the most noticeable feature of 
the series. At the same time the teeth are wide, the cement-areas 
broad, and the enamel simple, though at times broken up into a 
number of rings. So far as can be ascertained from the scanty 
amount of material the ridge-formula must have been low. 

Except with regard to the immense difference in size the 
characteristics of the molars described above, more especially in 
the lowness of the crowns ¢, appear to resemble more closely those 
of H. meridionalis than of any other of the larger Elephants of 
the Mediterranean Region. 

I]. Eneruas antiguus Falconer. 

As previously mentioned, remains of this Elephant $ had already 
been obtained from caves closeto Retymno. The teeth and bones 
noted below, and believed to belong to this species, were all pro- 
cured from a much damaged and fragmentary cave-deposit, one 
of several found close together in the cliffs bordering the south 
of Kharoumes Bay in the Eparkhia of Sitea|. Although evi- 
dently but a remnant of a formerly larger deposit, it was possible 

* Trans. Zool. Soc. vol. ix. p. 30, pl. 6. figs. 1, la. 

+ Leith Adams, Trans. Zool. Soc. vol. ix. pp. 108, 116. 

{ This characteristic of the molars of H. meridionalis was constantly noted by 
Falconer, see Pal. Mem. (London, 1868), vol. ii. pp. 128, 134, 138, &c. 

§ Signor Simonelli does not give the author of his H. priscus, but it may be sup- 
posed that this name is employed for the thick-ridged variety of EH. antiquus 
(Falconer’s HZ. priscus, Cat. Foss. Mamm. Brit. Mus. part iv. p. 122), since the name 
E. priscus of Goldfuss appears to have been applied to molars almost indistinguish- 
able from those of H. africanus (see Pomel, Bull. Soc. Géol. France, tome vii. 
1878, p. 51). 

|| Geol. Mag. n. s. dec. y. vol. i1. (1905), footnote 3, p. 199. 

244 MISS D. M. A. BATE ON [ Mar. 5, 

to trace the presence of Hlephant bones for a depth of several feet. 
Some few remains of ruminants, similar to those found in other 
parts of the island, also occurred here, but were only observed 
close to the uppermost of the bones of the Proboscidian, which 
probably became extinct long before these smaller mammals. 

Jt has already been mentioned elsewhere * that in another cave- 
deposit, on the same level—that is to say, not many feet above 
the sea, and only a few yards distant from the one under dis- 
cussion,—were found a number of land-shells, Helix pellita Fer., a 
species seemingly not previously recorded in a fossil state 7, these 
being preserved in the hard red breccia so common in cave-deposits. 
The presence of these shells points conclusively to the deposition 
of the mammalian remains under land-conditions, though it now 
appears just possible that these deposits were subsequently at all 
events partially submerged, which may help to account for their 
fragmentary condition. The occurrence of this movement is 
suggested by the discovery, in some sand adhering to a femur of 
HE. antiquus, of a large number of foraminifera and other marine 
forms, an account and list of which have been published by the 
Rev. R. Ashington Bullen=. Traces of a former submergence are 
more noticeable in the west of the island, especially at Sphinari 
and the Kutri and Haghios Basilis caves ; it was also unexpected 
in the east, where the coastal movement is supposed to have been 
for long past in an opposite direction to that in the west §. 

H. antiquus is represented by a number of limb-bones, including 
several perfectly preserved foot-bones, and a single right man- 
dibular ramus (M. 9384) containing twosomewhat damaged molars. 
Both from its size and from the fact of its having been found 
situated just above the limb-bones, it was at first sight thought 
that this last indicated the former occurrence in Crete of a small 
race of Elephant intermediate in size between the pigmy /. creticus 
and the very large species indicated by the remains noticed below. 
However, a further study of the material has caused the conclusion 
to be reached that it is a portion of the mandibular ramus of an 
immature specimen of #. antiguas, and that the teeth must be the 
penultimate and ultimate milk-molars, or perhaps the last of the 
milk and first of the permanent series. The general appearance 
and characters of these molars support this view of their identity, 
which is further strengthened by the fact of the large limb-bones 
occurring in the same deposit, and also that H. antiqwus has already 
been recorded from another district of the island. 

Lower Molars.—As already remarked, the two lower molars 
obtained, and believed to be those of #. antiquus, are situated in 
a portion of the right mandibular ramus shown in text-fig. 83 
and Pl. XIII. fig. 3: neither of these teeth is quite complete. 

* Geol. Mag. n. s. dec. v. vol. ii. 1905, footnote 2, p. 199; and Rev. R. Ashington 
Builen, Proc. Malacol. Soc. vol. vi., Sept. 1905, p. 307. 

+ Ibid. 

£ Geol. Mag. n. s. dec. v. vol. iii. pp. 353-358, pls. 18 & 19. 

§ For references to this, see Geol. Mag. n. s. dec. v. vol. ii. (1905), footnotes 2 & 3, 
10s IZ, 


The whole specimen is about 263 mm. (nearly 103 inches) in 
length ; the ramus is robust, the greatest thickness in the portion 
preserved being 112 mm., and its depth, measured in front of the 
first molar, is about 126 mm. Its anterior border is very abrupt 
owing to the advanced state of wear and consequent forward 
position of the anterior of the two teeth. 

Unfortunately neither of these molars retains its full com- 
plement of plates, which makes it impossible to determine with 
absolute certainty their exact position in the series, for, as may be 
seen by the specimens in the British Museum Collection, and has 
been pointed out by Dr. Leith Adams*, the milk-teeth of this 
species varied in size to a very great extent. However, it seems 
certain that the specimens in question represent either the two 
last milk-molars or the posterior of these and the first true molar. 

Text-fig. 83. 

Right mandibular ramus of Hlephas antiquus (2) bearing two lower molars. 
1 = . 
3 hat. size? 

The actual height of the foremost tooth (Pl. XIII. fig. 3 and 
text-fig. 83) is not shown, owing to the enveloping ramus; it 
consists of eight plates, but is extensively worn and _ projects 
considerably beyond the edge of the alveolus, so it is likely thate 
there may have been one, perhaps two, additional plates originally. 
The present length of the crown, measuring along the median 
line of the plates, is 92 mm., and its greatest width, which occurs 
at the sixth of the plates present, is 47 mm., its height above the 
ramus at the same place is 30 mm. ‘The cement-areas are not so 
wide as those of the following molar, but this is probably due to 
difference in‘vear ; in neither is there any mesial expansion of the 
plates. The enamel bands are rather thin and “‘ wavy” in outline, 
in both these respects differing from the specimens from Cape 

The second of the two teeth also shows eight plates, though it 

* Mon. Brit. Fossil Elephants, Pal. Soc. London, 1877. 

246 MISS D, M. A. BATE ON [ Mar. 5, 

is evident that one or more may be wanting posteriorly. Only the 
anterior five are worn, and the enamel is seen to be less irregular 
than in the preceding molar, considerable digitation of the plates 
is observable, while the cement-areas are of considerable width. 
This molar is remarkable for its very great height, being 103 mm., 
while the greatest width of the crown at the second plate is only 
43 mm. It will be remembered that this was given by Falconer * 
as one of the distinguishing characteristics of the molars of 
E. antiquus :—*‘ Great height of the plates. The height is more 
than double the width of the crown.” This is in striking contrast 
to the proportions of the teeth of Z. creticus, in some examples of 
which the height of the crown is exceeded by its width. Particu- 
larly in the rather thin and very wavy outline of the enamel bands, 
these specimens from the Kharoumes deposit resemble a number 
of those of H. mnaidriensis figured by Dr. Leith Adams 7, who 
also called attention to the resemblance between the molars of this 
Maltese species and those of 1. antiguas £. 

Limb-bones.—The specific or peculiar characters of the limk 
bones of #. antiquus do not appear to be well known, partly no 
doubt owing to the difticulty of distinguishing them in cases where 
the remains of more than one species of similar size occur in a 
single deposit, so that it is evidently chiefly by inference that the 
fragmentary collection of bones under discussion must be deter- 
mined as those of this species. However, it will have to be 
acknowledged that this contention is a strong one when we con- 
sider the identity of the teeth found in the same deposit, and the 
discovery of the cave near Retymno from which were obtained 
similar, though more complete, remains, At least two individuals 
are represented amongst the limb-bones, which number about 
twenty and are almost all imperfect, with the exception of a few 
foot-bones ; and in many cases the articnlar surfaces are damaged 
or missing, which makes it difficult to discern any features other 
than that of size. Nearly every specimen was covered with a thin 
red stalagmitic encrustation. 

The only portion of the spinal column procured is a portion of 
the neural arch of a dorsal vertebra (M. 9388). The collection 
contains two wlae (M. 9385), though only the proximal portion of 
each is preserved. One of these, which is unfortunately much 
crushed, is of the right side and apparently that of an adult. It 
seems to agree in size and general appearance with a corresponding 
bone of #. antiquus in the British Museum Collection (45203) §. 
A fragment of the humerus is still attached to the second specimen, 
which belongs to the left side and is that of a young individual, 
the line of junction between the shaft and the olecranon epiphysis 
being very apparent. On comparing it with the proximal portion 
of a left ulna in the Collection of the British Museum (45202) |j, 

* Pal. Mem. vol. ii. p. 176. 

+ Trans. Zool. Soc. vol. 1x. 

+ Mon. Brit. Fossil Elephants, Pal. Soc. 1877, pp. 25 & 50. 

§ Ibid. p. 59, D. 12. || Lbid. 


it appears that the depression below and between the olecranon 
and the projection of the sigmoid notch is more extensive in the 
former, and also that the distance between the coronoid process 
and the olecranon is comparatively greater. 

An imperfectly preserved left femur, with its distal extremity 
missing, was obtained, also the head of another and a fragment of 
the head of a third. The entire length of the former is 29 inches 
(89 mm.); it is decidedly short in the neck and has a shallow 
digital pit, both of which characters are claimed for #. antiquus by 
Dr. Leith Adams*, as well as for ZH. namadicus t+, E. africanus, 
and #. mnaidriensis. 

A left cuneiform, which is attached to the unciform, is very 
much larger than a corresponding bone in the British Museum 
Collection (36608) +, which, however, is by no means a large 
example for this species. The greatest width of the Cretan 
specimen, from its inner border to the outer angle, is 165 mm., 
while antero-posteriorly it measures 115 mm., though this would 
be slightly more were it quite intact. The height of its 
anterior face is 57 mm. The greatest height of the anterior 
face of the above-mentioned wnciform is 81 mm., antero- 
posteriorly it is 125 mm., and from side to side 151 mm. 
A considerably damaged right unciform appears to be of similar 

An almost perfect right trapeziwm included in the collection 
is found to differ considerably in size and outline from the one 
figured by Dr. Leith Adams, and considered by him to present 
diagnostic characters §. The former is altogether much smaller 
and slighter than the latter, its greatest height, taken perpen- 
dicularly, being 79 mm., and its circumference just below the 
external articular surface 180 mm. In the British Museum 
specimen the lower articular surface || is expanded and almost oval 
in shape, whilst in the one from Crete this is narrower and some- 
what elongated. The projection beyond the anterior border of the 
distal articular surface is extremely pronounced, and it is chiefly 
to this that the bone owes its irregular outline. 

The metacarpals ave represented by two distal extremities 
and an almost perfect specimen of the fifth metacarpal of the lett 
manus. Its exterior border is 142 mm., the proximal articular 
surfaces measure 83 mm. from inner to outer edge, while antero- 
posteriorly they are about 78 mm., the circumference at the centre 
of the shaft is approximately 210 mm. 

The proximal phalanx of the fifth digit of the right manus, 
another digital phalanx, an imperfect right semilunar, and some 
portions of ribs, are the only other well-preserved remains of 
EL. antiquus obtained. 

* Mon. British Fossil Elephants, p. 63. 

+ Considered by some writers as a variety of or identical with EH. antiquus (see 
G. E. Pilgrim, Records Geol. Survey India, vol. xxxii. part 3, pp. 204, 205, 215, &e.). 
+ Cat. Foss. Mamm, Brit. Mus. part iv. p. 136. ; 

§ Op. cit. pp. 160 & 234, pl. 19. figs. 9-9 B, B.M. 20821. 
|| Led. fig. 9B. 

248 MISS D. M. A. BATE ON [ Mar. 5, 

TIL. ConcLustons. 

It was at one time thought that the discovery of remains of 
different Elephants in the Pleistocene cave- deposits of Crete might 
help to throw additional, or perhaps even fresh, light on the subject 
of the origin and development of pigmy forms. However, it will 
have been seen from the above descriptions that the material ob- 
tained is very fragmentary, so that little more than the identity 
of the species can be established, though there is at least sufficient 
to suggest one or two minor problems of some considerable interest. 

The occurrence of two species of Elephant in more or less con- 
temporary deposits in such a comparatively restricted area as Crete 
hints at, though by no means proves, the likelihood of some 
possible relationship or connection between them. It seems quite 
certain that Crete, evenif well covered with a luxuriant vegetation, 
could not support for long an Elephant of such great size as 
Fi. antiquus, so it 1s probable that this large race, at all events as 
such, ceased to exist in early Pleistocene times, which would be 
tantamount to saying shortly after the separation of the island 
from the mainland. 

It seems a firmly established and well-supported theory that 
the dwarf proportions of the pigmy species found in most of the 
larger Mediterranean islands are a specialised character acquired to 
meet the exigencies of these island habitats ; for, as M. Gaudry has 
remarked *, these former small races have confirmed the idea that 
the size of animals was in accordance with the extent of their 
habitat. Among other arguments, if needed, in favour of this is the 
fact that up to the present no similar remains of a small form are 
known to occur in deposits earlier than Pleistocene. In this con- 
nection further information with regard to the extent and cause 
of the dwarfing of H. africanus pumilio Noack 7, from the French 
Congo, will be of the utmost interest, and might also help to 
elucidate the problem of the possibly analogous Hippopotamus 

Nowadays many writers maintain that #. antiqus is the parent 
form of the hitherto described pigmies of the Mediterranean aren, 
and this seems to have been well established by Prof. Pohlig = in 
the case of the remains from Sicily. At first sight the idea that 
these tiny forms were all derived from one of the largest species 
of the genus appears to be a somewhat startling one, but may it 
not be that the attainment of such dimensions under favourable cir- 
cumstances points to the species having been one highly susceptible 
to and strongly acted upon by its environment and conditions of 
life? If so, the same would apply under opposite ee 
adverse circumstances causing a rapid diminution in size; for, a 

* “Ont confirmé Vidée que la taille des animaux était en rapport avec l’étendue 
de leur habitat” ‘Foss. de Patagonie, Ann. de Paléontologie, tome i. fase. 11., Juin 
1906, p. 8. 

+ Zool. Anzeiger, Bd. xxix. No. 20, Jan. 1906. 

ft Abh. d. k.-bay. Akad. Wiss. xviii., and Nov. Act. L.-C. Akad. Naturforsch. lvii. 


Mr. Pilgrim has remarked*, it is to this adaptability that 
E. antiquus owed its wide geographical distribution and also its 
continued existence through a long period of time. 

Therefore, allowing that ZH. creticus is a dwarfed species, the 
question of descent is decidedly the most interesting point raised 
by this newly discovered pigmy of Crete. Two alternative theories 
with regard to this subject suggest themselves :-— 

A. That, taking into consideration the several points of resem- 
blance, which may prove to be only superficial, between the 
molars of ZL. creticus and HL. meridionalis, the latter may be 
the parent form, in which case it must have inhabited Crete 
contemporaneously with #. antiquus. 

B. That the Cretan race is descended from Z. antiquus, which 
we think is most likely to prove the true solution of the 
question. Some few points which appear to support this 
may be noted here. Among these are the fact of remains 
of this large species being found in the island and the 
evidence which tends to show a similar ancestry for other 
island races of the same region. 

Supposing #. creticus to be derived from Z. antiquus, this would 
necessarily mean that the former became differentiated since early 
Pleistocene times. That this calculation would allow ample time 
for such a change to be accomplished seems probable, when it is 
considered that /. eypriotes was the result of isolation for a no 
longer period, since, according to Messrs. Bellamy and Jukes- 
Browne *, it seems that a land-connection existed between Cyprus 
and the mainland as late as the beginning of the Pleistocene era. 

Both from the geological evidence and from the apparently still 
plastic condition of the small Elephants of Malta, as well as of the 
Hippopotamus of Sicily, at the time of their extinction, it is likely 
that these islands, more particularly the latter, have formed areas 
of isolation for an even shorter term than Cyprus. Therefore it 
would have been vital to the continued existence of the species 
that the dwarfing should be accomplished as rapidly as possible ; 
this also applies to the molars, in which the result might have 
been attained by diminishing the height and number of the plates, 
thus apparently to a certain extent reverting to a more simple 
condition. A significant fact which may lend weight to this sug- 
gestion is that, although #. mnaidriensis and #. melitensis of 
Malta are seemingly closely connected and probably represent two 
stages in the evolution of a pigmy from a large Elephant, yet the 
latter has the lower ridge-formula =, although it is the smaller and 
therefore, in that direction, the more specialised of the two. 

My sincere thanks are due to Dr. A, Smith Woodward, F.R.S., 

* Records Geol. Survey India, vol. xxxii. pt. 3, p. 216. 

+ ©The Geology of Cyprus, chap. viii. (Plymouth, 1905); and Bellamy, Key to 
Geol. Map Cyprus, pp. 15, 16 (Stanford, 1905). 

t Lydekker, Cat. Foss. Mamm. Brit. Mus. part iv. p. 151. 

for giving me every facility for working in his department of the 
British Museum (Nat. Hist.). 


Puate XII. 
Fig. 1. Inner side of the posterior part of the second and anterior part of the third 
right lower molars of Hlephas creticus. (M. 9383a.) | nat. size. 
2. Unworn plates of a molar of H. creticus. (M. 9379.) Nat. size. 
3. Crown view of right lower third molar of 4. ereticus. (M. 9381.) nat. 
Prats XIII. 
Fig. 1. Crown and (1 a) side views of second lower molar of H. creticus. (M. 9378.) 

+ nat. size. 
2. Crown view of second upper molar of #. ereticus. One of the plates has 

been restored. (M. 9377.) } nat. size. 
3. Crown view of the first and second right lower molars (or last milk-molar and 
first molar) of Hlephas antiquus Falconer. (M. 9384.) 3 nat. size. 

(The numbers of the specimens are those in the British Museum register.) 

3. Zoological Results of the Third Tanganyika Expedition, 
conducted by Dr. W. A. Cunnington, 1904—-1905.— 
Report on the Polyzoa. By Cuartes I. RoussELet, 

TReceived February 2, 1907. ] 
(Plates XIV. & XV.*) 

The freshwater Polyzoa collected in Lake Tanganyika by 
Dr. W. A. Cunnington are attached to stones and shells which 
were partly obtained in shallow water and partly dredged from 
20 to 40 fathoms. 

Altogether the collection contains five species, three of which 
belong ‘to the Phylactolemata and two to the Gymnolemata. 
Amongst the latter is Moore’s Arachnoidia ray-lankestert (107), 
which was found in some abundance on shells of Paramelania 
dredged from deep water. 

Two species of the Phylactolemata are of the Plumatella type, 
with horseshoe-shaped lophophore. One of these appears to be a 
new species, very closely adherent to stones, with half-formed 
ea-shaped tubes, which I have named Plumatella tanganyike. 

To the second species I have given no name, as the few frag- 
ments of tubes and the total absence of statoblasts offer no 
characters that would distinguish it from Plumatella repens. 

The third Phylactolematous specimen is an interesting new 
species of the genus Fredericella, which I have named Fredericella 
cunningtont in honour of its discoverer, who dredged it from 
25 fathoms near Mshale. 

The second of the Gymnolematous species is of special interest, 
as being always found associated with, and imbedded in, a fresh- 

* Wor explanation of the Plates, see p. 257. 
+ The numbers refer to the Bibliography, pp. 256, 257. 


Fig. FRDixon-Nuttall. del. Highley, del et lith. 



lee youll SOV Nell ONE 

Highley, del et lith. 

Fig 10. FER Dixon-Nuttali.del. 



water sponge, Spongila tanganyike Evans, only the small heads 
projecting beyond the surface of the sponge. Its affinities appear 
to approach nearest to Victorella, and I have therefore named it 
Victorella symbiotica. 

The following is a list of the five species here described :— 
Plumatella tanganyike, sp. n. 

. repens, var, ? 
Fredericella cunningtoni, sp. n. 
Arachnoidia ray-lankesteri Moore. 
Victoreila symbiotica, sp. n. 

The number of known species of Polyzoa inhabiting African 
fresh waters is thus brought up to eight, truly a remarkably 
small number for this vast continent. 

The other African species so far recorded are the following :— 

Fredericella sultana, found by Dr. Stuhlmann in Alexandria, 
Kgypt (7), and in the Equefa River, Natal, by the 
Hon. Thos. Kirkman *, 

Plumatella repens, var., from Ugogo, Victoria Nyanza, Albert 
Edward Nyanza,and Albert N yanza : the statoblasts only 
found by Dr. Stuhlmann (7) and also by Dr. Meissner 
on shells of Wtheria (in the Berlin Museum) from the 
Upper Nile, White Nile, the Niger, and Senegal (8, 9). 

Lophopodella (Pectinatella) carteri Hyatt, from Ugogo: 
statoblasts found by Dr. Stuhlmann (7). 

Lophopodella thomasi Rouss., from Hunyani River, Rho- 
desia (12). 

The indifferent preservation of the Tanganyika specimens has 
unfortunately hampered and prevented a complete study of all 
the species. They were narcotised with cocaine and then pre- 
served in alcohol, which is not a sufficiently good fixative for 
these animals. Freshwater Polyzoa must not be allowed to die 
in the anesthetic, which quickly macerates their delicate bodies. 
After treatment with cocaine in perfectly clean water they should 
be killed and fixed whilst still living, either with very weak 
osmic acid (;'5 per cent.) for ten minutes to half an hour, according 
to the age and actual strength of the solution, then washed, and 
also preserved, in 3 per cent. commercial formalin (97 c.c. water, 
3 c.c. formalin); or else fixed with a 10 per cent. solution of 
formalin (90 ¢.c. water, 10 c.c. formalin) for 24 hours, then 
preserved in 3 per cent. formalin to which 3 per cent. glycerine 
may be added to render the animals more transparent. Polyzoa 
fixed with osmic acid are stained brown, or even black, if allowed 
to stay too long in the fixative; those fixed with formalin remain 
white and transparent. 

The little bottles in which the preserved specimens are stored 
should have no air-space; an air-bubble plays havoc with the 
delicate tentacles of extended polypides. 

SUB Oo bo 

* Mentioned in his paper on the Rotifera of Natal as supporting tubes of Limnias 
ceratophylli: Journ. Roy. Micr. Soc. 1901, p. 232. 


With regard to the question of the origin of the freshwater 
Polyzoa, I am inclined to agree with Dr. Wesenberg-Lund (18), 
who has expressed the view that all the different groups have 
wandered from the sea, the Phylactolemata at an early period, so 
that their marine ancestors are not now known, and may have 
become extinct in the sea long ages ago, whilst the freshwater 
Gymnolemata, i.e. Paludicella, Victorella, Pottsiella, Arachnowdia, 
have immigrated at a much later period, so that they still show 
some aftinity with several marine genera. 

Tf this conception be correct, it follows that there can be no 
relationship between the living Phylactolemata and Gymnolemata, 
and still less can there be intermediate forms connecting the one 
with the other. 

In June of last year I succeeded, with the kind assistance of 
Dr. Bousfield, in again finding Victorella pavida in the Surrey 
Canal, London, where it had been obtained some twenty years 
ago, but not recorded since. A study of this species and of the 
other known freshwater Gymnolemata has impressed upon me 
that, in addition to the absence of statoblasts, they are all pos- 
sessed of a common character of considerable importance, which 
they share with a group of marine Ctenostomata, and which may 
well denote a certain degree of affinity. In all these forms there 
is a stolon which expands into a cell or zocecium, at the upper end 
of which an orifice is formed which may become enlarged into an 
elongated, more or less cylindrical tube for the protrusion of the 
polypide. Behind the orifice the stolon, after forming a septum, 
continues to grow out to form another cell a little further on ; 
then on each side of the cell normally one additional stolon arises, 
also separated by a septum, to form new branches, which repeat 
the same process of cell-formation until the growing point of 
the stolon is either broken off or becomes atrophied. In this way 
a zoarium is produced, forming an irregular network of branches, 
approximately at right angles to each other. This cruciform 
mode of growth can readily be observed in all the species of the 
following genera, which might be grouped together under the 
name of ‘ Cruciform Stolonifera ” :— 



Arachnidium. | nee 
oe 3 + Marine *. 
Cylindreciun, | 

PLUMATELLA TANGANYIK#, sp.n. (Plate XIV. figs. 1—4.) 

Specijic Characters—Zoarium consisting of clear light brown 
chitinous tubes, branching, curving, and interlacing, closely 

* The marine species Barentsia misakiensis from Japan, described by Dr. A. Oka 
in 1895, shows the same fundamental structure. 


adherent, encrusting stones, in the substance of which they are 
partly embedded ; also encrusting shells of molluscs; tubes some- 
times flat-sided. Tentacles about 20. Sessile statoblasts oval ; 
floating statoblasts not observed. 

The appearance of this new Plwmatella is very different from 
the European species and varieties of the genus, as will be seen 
by figs. 1 & 2, Pl. XIV. 

The tubes form a thin, closely adherent, interlacing, encrusting 
layer, and are partly embedded in the stone on which they grow ; 
or, possibly, stony crystalline material has been deposited in 
between and over the tubes. The tubes encrusting stones are 
fairly tubular, but those encrusting molluscan shells are flat-sided 
or e&-shaped in section, and the adhering side has only a very 
thin layer of chitinous material, so that the supporting shell is 
used by the animal to form part of its protecting tube. The 
tubes are of a light brown colour, clear and nearly transparent. 
The raised ends of the tubes are always tubular and white. 
Septa are present in the tubes at the poimts of branching. 

A full and well-extended head was not found; but there is 
every appearance that the lophophore is horseshoe-shaped, with 
about 20 tentacles. 

Sessile statoblasts were found in the tubes (Pl. XIV. figs. 3 & 4); 
they are oval in shape, but varying a good deal in their pro- 
portions of width to length, smooth, and surrounded by a thin, 
narrow, flat annulus without air-cells; their greatest size is: 
length “450 ue (iy in.), width 343 Ge in.). Ordinary floating 
statoblasts with swimming annulus were not observed. 

Diameter of tubes 321 y (5 im.) on average. 

Habitat. Encrusting stones just below water-level at Kalambo 
and Kassangiu; and also encrusting shells in shallow water at 



This specimen was found attached to shells of living 7iphobia, 
dredged from about 40 fathoms at Maswa. 

The tubes are brown in colour, semi-opaque, and show a 
deposit of fine material in more or less regular longitudinal or 
transverse lines; they are partly adherent, branching irregularly, 
tubular, and altogether have much resemblance to those of 
Plumatella repens of our lakes and canals, from which it is hardly 
possible to differentiate them. 

Fully extended specimens were not present, but the lophophore 
appears to have about 22 to 24 tentacles. 

The diameter of the tubes is 364 jy (,5 in.) on average. 

Some sessile statoblasts found in the tubes are oval, smooth, 
with thin flat annulus without air-cells, 407  (;4 in.) in length 
and 278 (_; in.) im width. Floating statoblasts with cellular 
annulus were ‘not observed. 

The tubes are larger, and the sessile statoblasts smaller and 
more elongated, than in the preceding species. 

Proc. Zcou. Soc.—1907, No. X VITI. 18 


FREDERICELLA CUNNINGTONI, Sp. n. (Plate XV. figs. 9 & 10.) 

Specific Characters.—Zoarium consisting of creeping, closely 
adherent, branching and interlacing €&-shaped tubes, built of a 
very thin, transparent, chitinous, internal membrane, covered 
externally with coarse grains of sand; lophophore circular, with 
16 tentacles. 

The appearance of the tubes of this new species is such that they 
were at first passed over as apparently the tubes of some aquatic 
worm or insect larva, until a circular lophophore was observed 
protruding from one of the tubes, when a closer examination, 
and the removal of the adhering tubes, revealed an abnormal 

Though the indifferent preservation of the specimens did not 
allow me to ascertain with certainty whether an epistome is 
present or not, all the other characters are those of a phylacto- 
lematous Polyzoon; and as the lophophore is circular, the genus 
Fredericella naturally suggested itself. 

The tube is low and €-shaped in section, very closely adherent 
to the surface of shells and stones, branching and interlacing 
freely. The structure of the tube is very abnormal; it consists 
of a very thin, transparent, chitinous layer, to the external 
surface of which grains of sand of various sizes and colour are 
cemented, not unlike the tubes of some marine worms. The 
grains of sand are mostly angular white quartz, and, to a less 
extent, green olivine, interspersed with red and black grains, and 
occasionally some plant and shell fragments. A few specimens 
show tubes made of finer sand-grains. Only the raised ends of 
tubes are tubular, otherwise the whole of the creeping tube is 
@-shaped, the adhering surface being free from grains of sand. 
The cells of the polypides are long, continuous, and with a few 
incomplete septa here and there. The width of the tubes is 
346 uw (4 m.) on an average. 

The lophophore is circular and has 16 tentacles, but the rest 
of the anatomy could not be made out, owing to the bad state of 
preservation. No statoblasts were found in the tubes. 

Hab. Encrusting stones and Weothawma shells dredged in 
about 10 fathoms at Kibanga, and also shells dredged in about 
25 fathoms near Mshale. 

The genus /redericella has contained, so far, only one species, 
i. e. #. sultana of world-wide distribution, if we consider the 
three American forms /. regina, walkottti, and pulcherrima 
as synonyms of /. sultana, in accordance with the opinion of all 
recent authorities. /. duplessisi, found by Dr. F. A. Forel 
(2, 3, 4) living unattached in the soft mud at the bottom of the 
Lake of Geneva, also is but a /. swltana which has adapted itself 
to its environment. Having examined specimens dredged by 
Dr. Forel, consisting of short tubes with few branches, not 
exceeding half an inch in length, I can find no difference between 
them and the type species. J should like to mention here that I 
find the tubes of /’, sultana rarely cylindrical, as has usually been 


stated; much more frequently they are distinctly angular, and 
mostly triangular in shape. 

ARACHNOIDIA RAY-LANKESTERI Moore. (Plate XIV. figs. 5 & 6.) 

In his book ‘The Tanganyika Problem,’ 1903, Mr. J. E. 8. 
Moore (10) mentions on p. 295 a gymnolematous Polyzoon found 
by him on shells of Paramelania dredged from about 20 fathoms 
off the shore of the lake, to which he gave the above name, 
considering it allied to the marine genus Arachnidiwm. 

This interesting species has been found again by Dr. Cunnington, 
also on shells dredged from 20 to 25 fathoms near Mshale. The 
colonies form an encrusting network, consisting of ovoid or 
irregular, membranous, brownish cells, with toothed edges, closely 
adherent to the shell, and connected together by narrow, tubular, 
creeping stolons. The cells or zocecia are very flat and shallow, 
and have at their upper end a tall erect tube from which the 
polypides protrude. The lophophore is circular, without calyx, 
and has 16 narrow tentacles. A septum can be seen at the origin 
of each stolon. 

In retraction the polypide retreats down the erect tube into 
the cell, invaginating in the process the anterior thin-walled part 
of the tube, which is the tentacle-sheath or kamptoderm. All 
that can be perceived within the shallow cell is a band of retractor 
muscles, and a sac filled with granules, possibly the stomach. 

This species belongs to the group of Gymnolemata which I 
have called “Cruciform Stolonifera,” and certainly appears closely 
allied to the marine genus Arachnidium. 

Its mode of growth appears to be as follows:—A narrow, 
filiform, closely adherent stolon, following all the irregularities 
of the surface of the shell, expands to form an ovoid cell, the 
edges of which are more or less toothed, the better to fit the 
depressions and ridges of the shell-surface. At the upper end 
of the cell an orifice is formed which elongates into an erect tube, 
whilst behind the orifice the stolon, after forming a septum, 
continues to grow to form another cell at some distance further 
on. At an early stage of the formation of the cells, stolons can 
be seen growing out laterally, one on each side, rarely more, to 
form side branches, and thus a very irregular cruciform network 
is formed. The cells are always well separated from each other 
by a stolon, and when found agglomerated together, as in 
Mr. Moore’s rough sketch, it is the result of two or more colonies 
having grown over and in between each other. 

Of the polypide the indifferent preservation does not allow me 
to say more than that the lophophore has 16 narrow tentacles. 

The size of the largest cell seen is 792 4 (34 in.); width of 
stolon 69 (s+ > im.); length of the tube when the animal is 
protruded 970 (4; im.). 

VICTORELLA SYMBIOTICA, sp.n. (Plate XV. figs. 7 & 8.) 

Specific Characters—Zoarium a narrow, tubular stolon with 


slightly expanded portions at imtervals, from which a long erect 
tube rises; side branches arise at the base of the zoecia. 
Polypides with cireular lophophore of 8 tentacles. The whole 
goarium and zocecia embedded in Spongilla tanganyike, on the 
surface of which the polypides protrude. 

The freshwater sponges brought back by Dr. Cunnington 
from Lake Tanganyika include specimens of Spongilla tanganyike 
Eyans, in which was found embedded this gymnoleematous polyzoon 
which cannot be identified with any known species. _ Its affinities 
seem to lie between the genera Cylindracium, Pottsiella, and 
Victorella; but its habitat in freshwater and the number of 
tentacles possessed by the polypides, have decided me to place it 
in the last-named genus. 

Potts’s Paludicella erecta (11), afterwards renamed Potisiella 
erecta by Prof. Kraepelin (6), is the only other freshwater species 
which is known to penetrate encrusting sponges, but its circular 
lophophore has 20 tentacles. 

The meandering cylindrical stolons, glassy white when cleared 
from adhering sponge-fragments, form an entangled mass difficult 
to follow, and are entirely embedded in the substance of the 
sponge. Adjoining and crossing stolons and tubes adhere, but 
can be separated by tearing apart. No septa have been observed 
in the stolons. 

The upright tubes, together with the slight expansions of the 
stolon, form the cell or zocecia, and lateral branches, one on each 
side, were observed springing from the expanded stolon, but these 
side branches are not always present. 

The tubes are long, erect, single, slightly widened at the base, 
of glassy transparency, except the small portion above the surface, 
which is rendered more or less opaque by fine granules, and 
emerge on the surface of the sponge where the polypides expand 
their small circular lophophore of 8 tentacles. The length of the 
tubes in this species is no doubt due to the necessity of reaching 
to the surface of the sponge. In Victorella pavida of Kent (1, 5) 
the cell is formed by a very distinct expansion of the stolon and 
the tube arising therefrom is of moderate length. Length of 
erect tubes up to 1:7 mm. (;/5 in.). 

It has been a matter of considerable difficulty to dig the 
complete stolon with the tubes out of the substance of the sponge 
without breaking up these delicate structures, and the operation 
has been only partially successful. 

Hab. Growing within and through specimens of Spongilla 
tanganyike, which were found encrusting rocks in shallow water 
at Ghamkaluki, and also on shells dredged from about 20 fathoms 
near Mshale. 

1. Bousrretp, E. C.—The Victorella pavida of Saville Kent. 

Ann. Mag. Nat. Hist. ser. 5, vol. xvi. 1885, pp. 401-407, 
1, plate. 








Du-Puessis-Gouret, Dr. G.—Essai sur la faune profonde des 
lacs de la Suisse. Mém. Soc. Helvétique d. Sc. Nat., 
vol. xxix. Bryozoaires. Genéve, 1885. 

Foren, Dr. F. A~-Faune profonde des lacs Suisses. Meém. 
Soc. Helvétique d. Se. Nat., vol. xxix. Bryozoaires. 
Geneve, 1885. 

——. Le Léman. Lausanne, 1904, vol. iil. pp. 115-114, 300. 

Kent, W. 8.—On a new Polyzoon, Victorella pavida, from the 
Victoria Docks. Quart. Journ. Micr. Se. new ser. vol. x. 
1870, pp. 34-39, 1 plate. 

Krarpretin, Prof. Dr. Karu.—Die Deutschen Siisswasser- 
Bryozoen. Hamburg, 1887. 

. Metssner, M.—Moosthiere. Deutsch-Ost-Afrika, Band iv. 


Beitrag zur Kenntnis der geogr. Verbreitung der 

Bryozoengattung Plumatella in Afrika. Zool. Anz. no. 430, 


Weiterer Beitrag zur Kenntnis der geogr. Verbreit- 
ung der Siisswassergattung Plumatella. Zool. Anz. no. 531, 

Moore, J. EK. $.—The Tanganyika Problem. London, 1903. 
Arachnoidia, p. 295. 

Ports, Epwarp.—On Paludicella erecta. Proc. Ac. Nat. Se. 
Philadelphia, vol. 1. 1884, pp. 215-14. Reproduced also in 
Ann. Mag. Nat. Hist. ser. 5, vol. xiv. 1884, pp. 437-439. 

Rovussevet, Cuas. F.—On a new Freshwater Polyzoon from 
Rhodesia (Lophopodella thomas). Journ. Quekett Mier. 
Club, ser. 2, vol. ix., April 1904, pp. 45-56. 

WESENBERG-LuND, Dr. C.—Biologiske Studier over Fersk- 
vandsbryozoer. Kj@benhavn, 1896, 4 plates. Résumé en 

PrarEe XIV. 

. Plumatella tanganyike (p. 252). Portion of interlacing tubes. X 15. 

Plumatella tanganyike. Flat under surface of tubes detached from shell, 
showing septa. X Jd. 

. Plumatella tanganyike. Sessile statoblast, upper surface. X 50. 

. Plumatella tanganyire. The same, under surtace. 50. 

. Arachnoidia ray-lankesteri (p.255). Colony growing on shell. X 20. 

. Arachnoidia ray-lankesteri. Single cell and polypide. X 60. 

Qe poe 

PratTe XV. 

7. Victorella symbiotica (p. 255). Piece of sponge with polypides protruding 
on surface. X 20. 
8. Victorella symbiotica. Cells and stolon separated from sponge. 20. 
9. Fredericella cunningtoni (p. 254). Interlacing tubes adhering to shell 
surface. X 15. 
10. Fredericella cunningtoni. Portion of same with protruding polypide. 
x 56. 


4. Zoological Results of the Third Tanganyika Expedition, 
conducted by Dr. W. A. Cunnington, 1904-1905.— 
Report on the Brachyurous Crustacea. By Wui.tiam 
A. Counnineton, B.A, Phe D., F.Z:S: 

{Received March 5, 1907. | 

(Plates XVI. & XVII.* and Text-figure 84.) 

1. Lntroduction. 

Since the year 1896, when Mr. J. EH. 8S. Moore paid his first 
visit to Lake Tanganyika, our knowledge of the fauna of that 
lake, and of the interesting problems which are connected with it 7, 
has considerably advanced. The single representative of the 
Brachyura then collected was described somewhat later t, but 
already, in 1887, a description had been given by Milne-Edwards § 
of another form brought from the lake by Captain Joubert. The 
description of this last-mentioned type was unfortunately very 
incomplete, while at that time no male specimen had been 
obtained. During Mr. Moore’s second expedition, in 1899-1900, 
he succeeded in collecting further examples of both the species 
already known, and furnished male specimens of the form which 
Milne-Edwards had described from the female alone. Finally, a 
Third Tanganyika Expedition was dispatched in 1904, with the 
conduct of which I had the honour of being entrusted, and which 
has added yet again to our knowledge of the Brachyurous 
Crustacea of the lake. 

The very earliest facts which were learned respecting the fauna 
of Tanganyika seemed to show that the animals it contained 
were not merely of an unusual type, but were different from those 
living in the other big African lakes. It is an interesting fact 
that, with the lapse of time and a very great increase in our 
knowledge of the lakes of Central Africa, this dissimilarity of 
Tanganyika has been not only confirmed, but rendered more and 
more striking. Among other animal groups, that of the Brachyura 
affords an excellent example of this remarkable state of things ; 
so that a double purpose will be served by the description in this 
paper of Crabs from Nyasa, which lake has the normal charac- 
teristics of the African fresh-waters. 

The collection of Crabs made during this Expedition contains 
representatives of five species, of which two alone have been 
previously described. By the kindness of Dr. W. T. Calman, I 
have been permitted to examine and compare with the collection 
a large number of specimens of nearly allied forms, belonging to 

For explanation of the Plates, see p. 276. 

Moore, ‘ The Tanganyika Problem ’ (London, 1903). 
Cunnington, Proc. Zool. Soc. 1899, p. 697. 

Ann. Sci. Nat. 7° ser., Zool. t. iv. (1887) p. 146. 



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the British Museum *. Among others are some which are stated 
to have come from Nyasa and Tanganyika; and since no notice of 
these specimens appears to have been published, it seems desirable 
to consider them, also, in the present paper. The following is 
thus a complete list of the species at present known to occur in 
the lakes in question :— 

Potamon (Potamonautes) inflatus (H. Milne-Edwards). 

¥5 (Potamonautes) orbitospinus, sp. i. 
i" (Potamonautes) sp. ? 


Potamon (Potamonautes) platynotus, sp. n. 
“ (Potamonautes) sp. ¢ 
Platythelphusa armata A. Milne-Edwards. 
maculata (Cunnington). 
conculcata, sp. 0. 



2. Systematic Notes and Description of New Species. 

Subfamily Poramontn a. 

Lake Nyasa. 

Poramon (Poramonautes) tNFLATUS (H. Milne-Edwards). 

Potamon (Potamonautes) inflatus Rathbun, Nouv. Arch. Mus. 
Hist. Nat. Paris, 4° sér., vil. (1905) p: 174 (ubi synon.). 

A single fair-sized male specimen in the collection of the 
British Museum (Reg. No. appears to belong to this 
species. The only particulars given are :—“ Kavisonda, Nyasa. 
Presented by Dr. Ansorge.” 

fig. 1.) 

Description.—Carapace considerably flattened, antero-lateral 
margins arcuated and denticulate, extending laterally less than an 
orbit’s breadth beyond the external orbital angle. No definite 
epibranchial tooth, but the end of the post-frontal crest forming 
a distinct corner at that point. Regions and sutures on carapace 
moderately marked, lateral regions exhibiting a series of small 
slightly oblique granular ridges. Front rather deflexed, less than 
one-third the width of the carapace, and with its anterior margin 

* By the courtesy of the authorities of the Berlin Museum, co-types of some of 
the species described by Hilgendort from German Hast Africa were lent to the 
British Museum for the purpose of comparison with those discussed here. I am 
especially indebted to Dr. P. Pappenheim for detailed information regarding these 


sinuated. _Post-frontal crest prominent, almost straight, and 
extending to margins, with branches of mesogastric groove 
angulated. Orbits large, and eyes large, with stout peduncles. 
External orbital angle produced into prominent spine. Ischium 
of external maxillipeds showing longitudinal furrow, somewhat 
nearer to the inner edge. Chelipeds in both sexes subequal ; 
merus trigonous, with a series of small’ spines and distally a pro- 
minent spine on the anterior margin; carpus with two spines on 
inner margin, the posterior being the smaller. Fingers pointed 
and slightly hooked; teeth fairly uniform, of moderate size. 
Ambulatory legs long and slender, also somewhat compressed. 
Colour in life dull dark green, shading into dark purple; the most 
prominent parts of the front, the external orbital spines, antero- 
lateral margins, and portions of post-frontal crest outlined in 
white; legs dark purple. 
Dimensions as follows :— 

Adult male (largest specimen) : 

Length of carapace ............ 384 
Breadth of carapace ............ 56°9 
Fronto-orbital width ......... 36°7 
NAAUGUG, GL THROWING sun odeos somone oa 13°8 
Adult female : 
eneth of carapace 22.5)... 36°9 
Breadth of carapace ............ 53°9 
Fronto-orbital width ......... 34°5 
AY AUCHEOL GE THEOL ad cddscddcocsene st 14-2 
Remarks.—As there has recently been published a very com- 

prehensive revision of the Potamonide by Miss Rathbun %, it 
becomes desirable to take this asa basis, and correlate with it any 
newly described forms. Accordingly that portion of the key 
given by Miss Rathbun’? which is affected is restated below, but 
slightly modified to include this new species, while distinguishing 
it from johnston, the species close to which it finds a place. 

h'. Carapace broader; less than three-quarters as long as broad. 
Jj. Antero-lateral margins denticulated; external orbital angle 

produced into prominent spine saa pecans aeeodnG orbitospinus. 
j'. Antero-lateral margins granulated; external orbital angle 
forms low blunt process ..............0...000ceetereeeccceeeeeess es Johnstoni. 

Even a casual study of the species comprised in the genus Potamon 
renders it abundantly evident that in many cases they cannot be 
distinguished by very striking differences of character. It is 
therefore the more important to make clear the actual points of 
disagreement which have been thought in this case to justify the 
formation of a new species. PP. orbitospinus approaches most 

* Nouv. Arch. Mus. Hist. Nat. Paris, 4° sér., vi.—viii., 1904-1906. 
+ Op. cit. vii. p. 160. 

nearly to the forms P. hilgendorfi (Pfeffer) and P. johnstoni 
(Miers). In order to render a comparison more easy, the 
diverging characters of these three species are given in tabular 
form, while it may here be stated that P. orbitospinus is a species 
certainly more easily recognised than some of those which are 

allied to it. 

hilgendorfi (Pfetf.) 

(not hilgendorfi Hilg.). TALMOS POPS SDs Le 

johnstoni (Miers). 

Carapace somewhat flat- | Carapace considerably 
tened. | flattened. 

Antero-lateral margins | Antero-lateral margins 
granulated. denticulated. 

External orbital angle | External orbital angle pro- 

} . . 
rather acute. | duced into prominent 

Fronto-orbital width °71 | Fronto-orbital width *64 
that of carapace. | that of carapace. 

Ischium of external maxil- | Ischium of external maxil- 
lipeds without longi-| lipeds with longitudinal 

tudinal furrow. | furrow. 

Posterior carpal spine | Posterior carpal spine (of 
(of cheliped) little | cheliped) sharp and 
developed. | well developed. 

| Ambulatory legs long and 

Carapace somewhat flat- 



External orbital angle 
forms low blunt process. 

Fronto-orbital width °59 
that of carapace. 

Ischium of external naxil- 
lipeds with longitudinal 

Posterior carpal spine (of 
cheliped) little deve- 

Ambulatory legs short 
and stout. 

| slender. 

Occurrence.—N kata Bay, 23/6/04. Several specimens of medium 
size. The Crabs were captured and brought me by the natives, 
so that I have no direct evidence of their mode of life. They are 
said to live in the water and not on land, though they sometimes 
come on to the beach. 

Three small and immature specimens in the collection of the 
British Museum (Reg. No. Only the following 
particulars are given :—‘‘ Lake Nyasa, Coll. Miss M. Woodward, 
pres. by Miss Sophia McLaughlin.” 

One rather small male, no precise locality given, British 
Museum (Reg. No. ‘“‘Lake Nyasa. Received from 
Mr. Joseph A. Williams, Universities Mission, Likoma, Lake 

One large female specimen, British Museum 
Bay to Ruarwe, June 1896.” 
H. H. Johnston.” 

A single large male specimen, collected by Mr. J. E. 8. Moore 
during one of his expeditions to Tanganyika, but without further 

(Reg. No. 
“West coast of Lake Nyasa from Ukala (? Nkata) 
“A. Whyte collector—pres. by Sir 


Poramon (POTAMONAUTES) sp. @ 

Reference has already been made to the fact that it is often 
difficult to distinguish between closely similar species of this 
genus. Many of the characters enumerated by the systematic 
writers on this group, while in certain cases affording a ready 
means of separating species, are in other cases of little or no 
value. A careful consideration of many of the described species 
and the examination of an extensive series of specimens lead to 
the undoubted conclusion that some species have been established 
without sufficient justification. While a species or even a genus 
may be established on the strength of a single specimen, if its 
characters are sufticiently unusual, it is necessary to be very 
cautious in doing any thing of the kind within the limits of this 
subgenus (Potamonanutes) in particular. Yet there are several 
species of Potamonautes based upon single specimens, and even 
upon specimens which were recognised as being immature. We 
do not know a great deal about the modifications in form produced 
by advancing age, but we do know, from the examination of any 
extensive series, that they are considerable. The proportion 
of length of carapace to its breadth, with the relative proportions 
of the front, the orbits, and the fronto-orbital width—all of 
them characters employed by the systematist—are undoubtedly 
dependent upon the age and growth ‘of the individuals. 

But more than this. A natural hesitation to lay stress on 
every little difference in form is only emphasized by the result 
of Schenkel’s* investigation of the species Potamon (Potamo- 
nautes) celebensis de Man. He is able to distinguish some six 
local varieties, in addition to the forma typica of de Man. These 
differ from one another in colour, in the shape of legs and carapace, 
in the extent of the sculpturing r, and especially as regards the 
degree to which the carapace is vaulted and the br omental regions 
dilated. But, as Schenkel points out, these features are precisely 
those which would be affected by a difference of surroundings. 
He finds the Crabs with the flattest carapace come from mountain- 
streams, where of course the water is pure and well oxygenated. 
The converse holds equally good, the gill-chambers in other 
varieties being inflated in proportion to the sluggish or muddy 
nature of the river inhabited, and it is obvious that the sculpturing 
is In a great Measure an expression of the degree of inflation of 
the carapace. Again, it is pointed out that the amount of the 
food-supply must have a powerful influence on growth: thus with 
two Crabs of similar size, that from water poor in nourishment 
will be far older than the other, and so not strictly comparable 
with it. 

As Schenkel considered it desirable to retain a single species, 
but to constitute a number of varieties in this case, this discussion 
of the facts is very suggestive in a general sense. Under such 
circumstances, some observers would not even separate into 

* Verh. naturf. Ges. Basel, Bd. xiii. 1902, p. 528. 


varieties, and, on the other hand, there can be no doubt that many 
so-called species of other writers are not more distinct than these 
varieties of Schenkel. If it is possible to trace a gradual 
transition from one variety to another, as appears to be the case 
with the form celebensis, it becomes evident that certain established 
species are of similarly little value, and that species of this genus 
ought never to be constituted on such slender grounds. 

This matter is dealt with at such length, because the conviction 
has forced itself upon me that the number of African species, at 
all events, has been multiplied to an excessive degree. Where 
two or more well-marked species exist side by side, there can be 
no hesitation in the matter; but our knowledge of the Crabs of 
the African continent is still very scanty, and over a vast area 
restricted to a few isolated records. Thus it is impossible to 
determine yet whether this or that species may not prove to be only 
a slight local modification of some widely distributed form. This 
being the case, I have not ventured to name the two specimens 
from Nyasa, which are placed under the head-line above. 

Although it was during my study of the two species described 
as new in this paper that I became impressed with the unsatis- 
factory character of some species of Potamonautes, it was not 
until [ examined these two Nyasa specimens and a third from 
Tanganyika that [found a case where the most careful comparison. 
only resulted in confusion with several closely allied species. The 
two specimens from Nyasa—a rather small male and a small 
female—are in the collection of the British Museum (Reg. No., and the following note is given as to locality :— 
“ Kondowe to Karonga, N. Nyasa, June 1896. A. Whyte 
collector—pres. by Sir H. H. Johnston.” 

Using asa basis the key in Miss Rathbun’s recent revision, 
we find they belong to group c, where the lateral margins extend 
less than an orbit’s breadth beyond the external orbital angle. 
Further, they would seem to belong to division e’, where the 
post-frontal crest is little advanced in the middie, and to sub- 
division f', with fingers of the cheliped but slightly gaping. 
Thus they would come nearest to the species hilgendorfi ( Pfeff.) and 
johnstoni (Miers). The specimens differ from hilgendorfi (Pfett.) 
principally in possessing a furrow on the ischium. From johnstont 
they differ in being distinctly longer in proportion to the breadth, 
in the sculpturing being less marked, and in the absence of the 
fairly stout spinules on the anterior margin of the carpus of the 
cheliped—a single distal spine only being present. 

They agree to some extent also with suprasulcatus pseudo- 
perlatus, which, if it exist at all as a true variety, is In a Measure 
intermediate between hilgendorfi (Pfeft.) and johnstont. The 
length in proportion to the breadth is, however, more extreme in 
the case of suprasulcatus pseudoperlatus. From another important 
type in group c—perlatus itself—these individuals do not differ 
greatly, the carapace being of much the same proportions as 
regards length and breadth. The width of the front is, however, 


considerably less than in perlatus, and the post-frontal crest does 
not extend forward so far in the middle. The posterior carpal 
spine of the cheliped is also sharper and more prominent. Both 
the male and female specimens show only a single transverse 
furrow on the sternum in front of the abdomen, while in the 
specimens of allied species examined the males almost invariably 
have two furrows even in young individuals. 

From this description of the characters of these two Nyasa Crabs, 
it will be evident that my former contention is not without 
foundation. These individuals agree to a considerable extent with 
several species, and where they disagree with one they agree with 
another, while, to my mind, they have not a single character 
which satisfactorily distinguishes them from those forms to which 
they are undoubtedly allied. In the present state of our know- 
ledge, it would be futile to attempt to determine which species 
should be eliminated, and Miss Rathbun has certainly taken the 
vight course in retaining most of the species hitherto described, 
though it seems to have led her into some difficulties respecting 
her key. Bearing in mind the facts which have been detailed at 
some length above, I prefer, in a case like this, to leave the 
specimens unnamed, and so at least avoid adding to the already 
existing superfluity. 

Lake Tanganyika. 

figs. 1 & 3.) 

Description.—Carapace much flattened, antero-lateral margins 
aveuated and denticulate, extending laterally shghtly more than 
an orbit’s breadth beyond the external orbital angle. No definite 
epibranchial tooth, but the end of the post-frontal crest forming 
a distinet corner at that point. Regions and sutures on carapace 
ill-defined, postero-laterally a few inconspicuous granular ridges. 
Front rather deflexed, less than one-third the width of the cara- 
pace, and with its anterior margin sinuated. Post-frontal crest 
prominent, slightly arcuated, but sinuated laterally, extending 
to margins; branches of mesogastric groove straight. Orbits, 
eyes, and peduncles small. External orbital angle existing as a 
short blunt spine. Ischium of external maxillipeds not showing 
longitudinal furrow, though this may be faintly mdicated in the 
oldest specimens. Anterior portion of the sternum hairy in the 
female, not hairy in the male. Chelipeds in both sexes subequal ; 
merus trigonous, with a series of granules, and distally a stout 
spine, on the anterior margin; carpus with a prominent spine on 
the inner margin and a slight process just behind it. Fingers 
distally hollowed out, spoon-shape and meeting closely in a sharp 
cutting-edge (Plate XVII. fig. 3); dactylus usually slightly 
longer than the pollex. Teeth fairly uniform, of moderate size, 
but the fingers of the larger chela gape a little and have proximally 
a few larger flat crushing-teeth. Ambulatory legs of moderate 
length, little compressed. Colour in life a uniform dull greenish 
brown ; the dactyli of the chelipeds in the males usually black. 


Dimensions as follows :— 

Male (probably young specimen) : mm. 
Length of carapace............-.. 20°3 
Breadth of carapace ...........- 30°7 
Fronto-orbital width ............ IS 7 
ANVGWGKGli it MNROVMG “gececbeseeovscoco: 10-1 

Adult female (largest specimen) : 

Length of carapace ............ 33°7 
Breadth of carapace ...........- 48-2 
Fronto-orbital width............ 30-0 
VivatGlelhh Ohi TORO, Coo gecAeecooeBanns 13°83 

Remarks.—This species comes into Miss Rathbun’s key under 
the heading c', where the carapace extends laterally more than 
an orbit’s breadth beyond the external orbital angle. It finds its 
place in the same subdivision (g') as the species ambiguus and 
mrogoroensis, and that portion of the key, modified to include it, 
is accordingly given below : 

g'. Anterior branch of cervical suture absent. 
h. Ischium of external maxillipeds without longitudinal furrow. 

;, Anterior portion of female sternum hairy................... 0 | latynotus. 

3 JOM 
j’. Anterior portion of female sternum TOG |NEVVO cosees _... Immrogoroensis”™. 
7’. Ischium of external maxillipeds with longitudinal ftrrow... ambiguus. 

The principal points in which platynotus differs from ambiguus 
on the one hand and suwprasulcatus on the other are also given in 
tabular form. The most striking features of the new species are 

suprasulcatus Hilg. | platynotus, sp. Nn. ambiguus Rathbun. 

Carapace moderately vaulted. Carapace much flattened. Carapace distinctly vaulted) 

Antero-lateral margins granu- Antero-lateral margins Antero-lateral margins 
lated. _ denticulated. granulated. 

Anterior branch of cervical Anterior branch of cer- Anterior branch of cervi- 
suture present. vical suture absent. cal suture absent. 

Fronto-orbital width °60 that Fronto-orbital width °62 | Fronto-orbital width °57 

of carapace. | that of carapace. that of carapace. 

Ischium of external mavxilli- Ischium of external | Tschium of external maxil- 
peds without longitudinal maxillipeds without lipeds with longitudinal 
furrow. | longitudinal furrow. furrow. 


Fingers pointed and slightly Fingers distally hol- | Fingers pointed and 
hooked. lowed out spoon-shape. shghtly hooked. 

the flattened carapace, the denticulated antero-lateral margin, and 
the character of the fingers. 

* Tt is by no meaus obvious why Miss Rathbun has inserted the form mvrogoro- 
ensis here, under the heading c’, when it is quite clear from Hilgendort’s description, 


Occurrence.—Kasakalawe and Kituta Bay, both south end of 
Tanganyika. Several specimens; some females probably full- 
grown, but the largest male apparently not so. The Crabs were 
taken under boulders about high-water level. 

Poramon (PoTAMONAUTES) sp. 

Under this heading, I placea single rather small male belonging 
to the British Museum (Reg. No., of which the only 
particulars are: “ Lake Tanganyika, E. C. Hore (ex coll.).” This 
specimen agrees with and differs from its near allies in very much 
the same way as the unnamed forms from Nyasa, which are 
dealt with above. Thus it finds its place near to the latter, with 
which it agrees in showing only a single transverse furrow on the 
sternum in front of the abdomen, although a male. It disagrees 
with the Nyasa form, however, in being less inflated and in 
showing the sculpturing more distinctly. Further, the antero- 
lateral margins are more finely perlated, and the spime on the 
merus of the chelipeds is longer and sharper. In this case also 
the specimen differs little from several species, and appears to 
have no satisfactory distinguishing characters of its own, so that 
I again take the course least open to objection and leave it 

Genus PLuaryTtHEeLpHusSA A. Milne-Edwards. 

Platythelphusa A. Milne-Kdwards, Ann. Sci. Nat., Zool. 7° ser., 
t. iv. (1887) p. 146. 

Platytelphusa Hilgendorf, Deutsch-Ost-Afrika, Bd. iv. Lief. ix. 
(1898) p. 21. 

Limnothelphusa Cunnington, Proc. Zool. Soc. 1899, p. 698. 

Limnothelphusa and Platythelphusa Moore, ‘The Tanganyika 
Problem’ (London, 1903), pp. 280, 286. 

Platythelphusa and Limnothelphusa Rathbun, Nouv. Arch. 
Mus. Hist. Nat. Paris, 4¢ sér. vii. (1905) pp. 268, 269. 

There area number of reasons which have led me to offer here 
a new description of this genus established by A. Milne-Edwards in 
1887. AsI have stated elsewhere *, the account given of the type 
species, P. armata, and particularly the figures of it, leave a good 
deal to be desired. Miss Rathbun reproduces a photograph of 
this same individual, and adds something to the description; but, 
as a result of the Second and Third Tanganyika Expeditions, we 

very incomplete though it is, that his specimen should come into the group e¢, in 
which the carapace extends laterally less than an orbit’s breadth beyond the external 
orbital angle. While it may perhaps be doubted whether the distinction given 
above under j and j’ is of much weight, it will be evident that if the species 
mrogoroensis were withdrawn from its false position, the new form platynotus 
would then be distinguishable from ambiguus by the difference of the external 
maxillipeds. It is not necessary to discuss here the position which mzrogoroensis 
should really occupy, the simpler course is taken of merely incorporating the new 
species in the existing key. 
* Proc. Zool. Soc. 1899, p. 701. 


now possess in this country a fairly complete series of this species 
of both sexes. We have the possibility for the first time of com- 
paring males with the female on which the genus was founded, 
and this might in itself justify a redescription. Buta comparison 
between a large series of the form described as Limnothelphusa 
maculata and the specimens of Platythelphusa armata (which I 
was unable to make in 1899), has convinced me that the former 
cannot be regarded as constituting a separate genus, but falls mto 
place as a species of Platythelphusa. A third species of Platy- 
thelphusa is among the acquisitions of the last expedition, so that 
the description which follows has been materially modified in 
view of our much more extensive knowledge. 

Description.—Carapace almost quadrilateral ; antero-lateral 
margins arcuated and armed with spines ; postero-lateral margins 
but shightly arcuated., Front little deflexed, nearly straight. Post- 
frontal crest distinct and perlated, but not extending to lateral 
margins. Sub-orbital spine more or less distinct, in addition to 
prominent mner sub-ocular tooth. A stout triangular process 
descends from the external angle of the front, and may be pro- 
duced into a small spine antero-distally. Antenne situated partly 
behind and partly between this descending process of the front 
and the inner sub-ocular tooth ; the distal segments thus escape 
an appearance of displacement by the front. Merus of external 
maxillipeds broader than long, the palp being attached to its 
antero-internal angle; ischium without longitudinal furrow. 
Ambulatory legs somewhat compressed, the fourth leg consider- 
ably shorter than the others, and with its two terminal segments 
broad and flattened. 

Remarks.—With the inclusion of Limnothelphusa, this genus 
has lost nothing of its original distinctiveness. In the generic 
description of Limnothel, phust, stress was laid on the simple 1 nature 
of the second antennal segment, which was undistorted by the 
deflexed front. Although it was perhaps not very happily ex- 
pressed, this character is just as typical of the other two species 
we know from Tanganyika, and deserves emphasis accordingly. 
In those genera and subgenera where the front is considerably 
depressed, the antenne have the appearance, at least, of a lateral 
displacement or distortion in consequence. All the species of 
Platythelphusa show the front little deflexed, but have a trian- 
gular process descending from the external angle, which process, 
however, passes to the front and side of the antenna, without 
modifying its shape or direction. In order to make this distinctive 
feature quite clear, the frontal region of all three species of Platy- 
thelphusa is figured, while corresponding figures are given of 
certain well-known species of the subgenera of Potamon (Plate X VI. 
figs. 2-7). 

Owing to the modification of the generic characters of Platy- 
thelphusi and the suppression of ‘the genus Limnothelphusa, 
Miss Rathbun’s key to the subfamily Potamoninz * requires 

* Op. cit. t. vi. p. 245. 


alteration. In the restatement which follows, the subgenera are 
omitted, as they are not affected, and care has been taken to 
retain the original form as far as possible. 

Subfamily PoraMonin”. 

e. Ocular peduncles large, not tapering towards distal extremity. 
f. A stout triangular process descends from the external 

angle of the front ............. PLATYTHELPHUSA. 
S'. No process desuanding Bunn the axiom anala a the 
g: Front not armed with spines or spinules ................... Poramon. 
g'. Front armed with spinules ............ HyYDROTHELPHUSA. 

. Ocular peduncles small, tapering tow andes ictal: amenity, TERIMETOPUS. 

Since there are now three species included in the genus Platy- 
thelphusa, 1t may be well to furnish a key to them, although they 
are much more distinct and well defined than is the case with 
many species of Potamonautes. 

Key to the Species of PLAYYTHELPHUSA. 

a. Carapace extending laterally more than an orbit’s breadth beyond 
the external orbital angle; front less than one-third width of 
carapace .........- . armata. 
a’. Carapace exionitine TleGeonetlly: Tees ann an + ors s ineadihe he dnd 
the external orbital angle; front more than one-third width of 
b. Carapace moderately convex; carpus of cheliped without spines 
on outer margin; ambulatory legs of moderate length, anterior 
margin of the merus without spines ........ .. maculata. 
b'. Carapace extremely tlattened ; carpus of ghelipadl Theanine spines 
on outer margin; ambulatory legs long, anterior margin 
of the merus produced distally into two spines..................... conculeata. 

PLATYTHELPHUSA ARMATA A. Milne-Edwards. (Text-figure 84.) 

Platythelphusa armata A. Milne-Kdwards, Ann. Sci. Nat., 
Zool. 7° sérv., iv. (1887) p. 147. 

Platytelphusa armata Hilgendorf, Deutsch-Ost-Afrika, Bd. iv. 
Lief. ix. (1898) p. 22. 

Platythelphusa armata Moore, ‘The Tanganyika Problem’. 
(London, 1903), p. 286. 

Platythelphusa armata Rathbun, Nouv. Arch. Mus, Hist. Nat. 
Paris, 4° sér., vil. (1905) p. 269. 

Description.—Carapace moderately convex, extending laterally 
more than an orbit’s breadth beyond the external orbital angle. 
The antero-lateral margins differ little in length from the postero- 
lateral, with which they are almost directly continuous. Number 
of spies on antero- lateral mar ein extremely variable, usually 
four or five, in addition to the spine at the external orbital angle. 
Regions and sutures fairly well marked. The postero- lateral 
regions exhibit a series of small slightly oblique granular ridges. 


Front less than one-third the width of the carapace, its margin 
perlated, and with a stout spine at each extremity. Sub-orbital 
spines short and stout; a small spine on the descending process 
of the front. Orbits small, -16 width of carapace ; orbital margins 
perlated. Eyes and peduncles small. Chelipeds in both sexes 
unequal; merus trigonous, with distally a prominent spine on 
the anterior margin, and a small spine or tubercle on the ventral 
margin ; carpus with two spines on inner margin and one above 
the point of articulation with the hand. Hands and fingers 
somewhat compressed, the dorsal border being keeled and granu- 
lated ; the ventral border in the larger chela characteristically 
arcuated at the junction of the pollex. Fingers pointed and 
slightly hooked ; those of the larger chela gape, and have a pair 
of large flat crushing-teeth proximally, with a series of rather 
smaller ones distally. In the smaller chela the teeth are fairly 
uniform and of moderate size. Ambulatory legs of moderate 
length. Colour, in life, yellowish brown, with irregular blotches 
of darker brown on the carapace. 

Text-fig. 84. 

Platythelphusa armata, large male. X 3. 

Milne-Edwards and Miss Rathbun give the detailed dimensions 
of an adult female (type specimen), but as we now possess for 
the first time particulars of males, the measurements of an 
adult male—the largest known specimen—are included here. 
In this individual (text-fig. 84) the size of the larger chela is very 
striking, the length of the hand and pollex being greater than the 
whole breadth of the carapace. 

Proc. Zoot. Soc.—1907, No. XIX. 19 


Adult male (largest specimen), Moore’s collection : 

Wenethvoncanapace werner arrears eee re rae eC ere 53°2 
TereeAVGlilN Olt (CARGAORIE®, 9 Ghoasosv onvaddacboceubanvencoacades 66°3 
JASN H-COre OMA VAMGHEIO papaccoopeseasdosodonacconnponcoouse 40-1 
Waadth Of fromb Vili cores. een esc ae atone eee 18-7 
Larger chela: greatest length of hand and pollex. 71:3 

Pt ” greatest height of hand ............ 35°8 

Remarks.—While the chelipeds in both sexes are unequal, it 
appears that either the right or left may be the larger, quite 
indiscriminately. The large crushing-teeth are often considerably 
worn in old specimens, and the spines of carapace and chelipeds 
become blunt and rounded. ‘This is, in fact, the case with the 
male specimen which is figured, where it is clear that certain of 
the spines have lost their original sharpness, but it should be 
understood that the spines in question are normally very sharp 
and strong. ; 

A curious feature of the specimens of this Crab is the number 
of circular blotches which appear on the surface of the carapace 
and appendages. The marks are approximately round, and seem 
due to an eating-away of the calcareous matter of these spots, 
which occur in greater numbers on larger and older specimens. 
In some cases the fingers of the chele have suffered severely, 
being partially eaten through by this process of erosion. It 
seems most probable that these blotches are due to the action 
of boring Alge. A portion of test including such a spot was 
decalcified, cleared, and mounted in balsam, but showed little 
structure even then. From a central point, a number of fine 
processes, or perhaps tubules, could be seen to radiate, but nothing 
further could be made out. 

Occurrence.—The locality is given by Milne-Edwards simply 
as Lake Tanganyika. Moore states* that he obtained this species 
only off the west coast of the lake, and in nets and dredges worked 
in water of about 20 fathoms. The experience of the third 
Expedition shows that it occurs more widely distributed in the 
lake, and may be taken in much shallower water than 20 fathoms. 

Kasakalawe (south end): a single, rather young specimen, 
taken under a stone about water-level, at the same time as speci- 
mens of Potamon (Potamonautes) platynotus. 

Mbete (south end): two or three young specimens came on 
board my dhow clinging to the anchor-chain. The vessel was 
probably anchored in two or three fathoms of water. 

Among a number of specimens of Platythelphusa maculata, 
dredged in 10-15 fathoms, principally at the south end of the 
lake, are two or three small Crabs belonging to this species. 

Vua, on the west coast: a single adult, seen from the dhow 
crawling about in a foot or two of water, and caught in a hand- 

North end of the lake: a single specimen not full-grown. 

* Op. cit. p. 280. 


figs. 5 & 6.) 

Limnothelphusa maculata Cunnington, Proc. Zool. Soc. 1899, 
p. 698. 

Limnothelphusa maculata Moore, ‘ The Tanganyika Problem ’ 
(London, 1903), p. 280. 

Limnothelphusa maculata Rathbun, Nouv. Arch. Mus. Hist. 
Nat. Paris, 4° sér., vil. (1905) p. 269. 

As this species is now regarded as falling under the genus 
Platythelphusa, in consequence of the examination of much more 
extensive material, it becomes evident that it merits a more 
complete specific description, which accordingly follows. 

Description.—Carapace moderately convex, extending laterally 
less than an orbit’s breadth beyond the external orbital angle. 
Antero-lateral margins shorter than postero-lateral, with which 
they are almost directly continuous. Number of spines on antero- 
lateral margin variable, usually three, in addition to the spine at 
the external orbital angle. Regions and sutures moderately 
marked. Postero-lateral regions exhibit a series of small slightly 
oblique granular ridges. Front more than one-third the width of 
the carapace, margin perlated, and in extreme cases produced into 
a small spine at each extremity. Sub-orbital spines little deve- 
loped: no spine on the descending process of the front. Orbits 
large, -24 width of carapace; orbital margins perlated. Eyes and 
peduneles large. Chelipeds in the male unequal, subequal in the 
female; merus rather short, trigonous, with distally a sharp spine 
on the anterior margin ; carpus with two spines on inner margin 
and one above the point of articulation with the hand. Fingers 
of the larger chela gape considerably, are pointed and slightly 
hooked, and carry a few larger crushing-teeth proximally, with 
smaller teeth distally. Fingers of the small chela, and both chele 
of the female (Plate XVII. fig. 5), distally meet closely in a sharp 
cutting-edge, while proximally there are uniform teeth of moderate 
size. Ambulatory legs of moderate length. Colour in life dark 
brownish grey, with dark brown or reddish spots. 

As now there has been obtained quite an extensive series of 
specimens, some of which are larger than those examined before, 
it seems worth while to furnish a further list ef dimensions, to 
compare with those given under the original description of this 
species. As in the case of P. armata, the larger chela of old 
male specimens is of a great size, the length of hand and pollex 
exceeding the total breadth of the carapace. 

Adult male : mm. 
ene thgoiscanapace mee \seeseny ees eee ere eens 12:9 
iBveaddthpofrcarapacem ante cee oon eee 16-4 
Hhgorango=ore owe Ml WCU copencossodoseeanooodoveconbacanuror 13:2 
Ect v0 feito 1G yee VS 0 ac Mea he ere 6-1 
Larger chela: greatest length of hand and pollex 16:9 

Bs » greatest height of hand ............ 8:7 


Adult female (largest specimen) : 

Taength of carapace © .........cecmeceueescoeenecemen cs 13°1 
Brena tlaareCama Paces cacsegs-acine Sorevige arse ceataeeteee 16°9 
Hirombo-or Oita wid hit caren -akee reer eeer ach ene tere: 13°2 
WWadthvofsronk 261 atht-ereear fidabadopeaantinatetucs 6:1 

Remarks.—While none of the specimens examined in 1899 
carried ova, it is satisfactory, on comparison with this more ex- 
tensive collection, to find that those regarded as adult were so in 
reality. Female specimens of approximately the same size as that 
described, but one of which is ovigerous, while the other carries 
the already liberated young, occur in the recent collection. 
Indeed we have further a female with total breadth of only 
12:4 mm., but which is nevertheless ovigerous. The average 
size of the eggs themselves, which are not quite round, 1s 
IS) Se Ihss) iar, 

The much smaller size of this species enables it to be distin- 
guished at once from adult specimens of P. armata, but, apart 
from that, the great relative breadth of the front and size of the 
orbits are differences easily recognised. Among the number of 
specimens which we now possess there exists considerable indi- 
vidual variation in respect to the development of certain spines, 
and this accounts for a slight discrepancy which may be noticed 
between the foregoing description and that given in the first 
instance. In the original type specimens, the spine on the carpus 
of the cheliped, above the articulation with the hand, is indicated 
but slightly, if at all. As it is, however, very well developed in a 
number of individuals, it is now included among the specific 
characters. Again, certain of the new specimens exhibit a small 
spine at each extremity of the front, and have a slight indication 
of sub-orbital spines, while others show no trace of these features. 
A more complete description of the fingers of the chelipeds is 
now given, and from my own observations I can add the colora- 
tion during life. 

Occurrence. —The type specimens described in 1899 were 
obtained, according to the information supplied me by Mr, Moore, 
from Kituta Bay (south end), while he had also taken specimens 
at Niamkolo (south end) and Sumbu on the west coast. The 
Crabs were said to have been captured in fairly deep water—never 
less than 60, and from that to 500 feet deep. In his book on 
Tanganyika, however, Moore states* that the specimens were 
obtained in water varying in depth from 500 to 600 feet. He 
also adds that the Crab occurs throughout the lake. With the 
latter statement, the experience of the recent Expedition is quite 
in accord; but whether the original individuals came from such 
very deep water or not, it is certainly a fact that the species may 
be found in much shallower regions. 

Niamkolo Bay (south end): a large number of specimens 

* Op. cit. p. 280. 


dredged among shells in about 10-15 fathoms. Also obtained in 

rather shallower water, but seldom in less than about 8 fathoms. 
Pembe (east coast): among shells dredged in about 10 fathoms. 
Kirando (east coast), 1/12/04: dredged in about 10 fathoms. 

Among these, a curiously blotched and not spotted individual. 

PLATYTHELPHUSA CONCULCATA, sp. n. (Plate XVII. figs. 2 & 4.) 

Description.—Carapace extremely flattened, extending laterally 
less than an orbit’s breadth beyond the external orbital angle. 
The whole body remarkably thin in appearance. Antero-lateral 
margins shorter than postero-lateral, with which they are almost 
directly continuous. ‘Three spines on antero-lateral margin, in 
addition to that at the external orbitalangle. Regions and sutures 
very ill-defined. Lateral regions exhibit a few small granulay 
ridges. Front more than one-third the width of the carapace, 
margin spinuliferous and with a sharp spine at each extremity. 
Sub-orbital spines prominent; a small spine on the descending 
process of the front. Orbits of moderate size, 18 width of cara- 
pace; orbital margins perlated. Eyes and peduncles rather large. 
Chelipeds in the male subequal ; merus trigonous, with distally a 
sharp spine on the anterior margin, and a small blunt spine on 
the ventral margin; carpus with two spines on inner margin, two 
on outer margin, and one above the point of articulation with the 
hand. Fingers distally meet closely in a sharp cutting-edge, 
while proximally there are uniform teeth of moderate size. 
Fingers of the larger chela (Plate XVII. fig. 4) have a few larger 
teeth proximally. Ambulatory legs long and slender ; the anterior 
margin of the merus, in each case, produced distally into two 
sharp spines, of which the terminal one is inconspicuous on the 
first and fourth legs. Colour (in spirit) yellowish brown, with 
darker reddish-brown spots. 

Dimensions as follows :-— 

Male (probably adult) : fai 
Meneth of carapace ..\sheass sn naae- 2-00 ne 10:0 
Breadt ln olveaid PRE: ./Mas ce dastemal- oleae a> e9 
Bvouto-oroibaly WiGtle i -l)jasety dae) eet ele ol 
WW icitherotficonit 271), gue seas aaare tote. ade 50 

Remarks.—It is unfortunate that we possess only a single 
specimen of this species, which appears nevertheless to be quite 
well marked. In size it is still smaller than P. maculata, while 
the much flattened carapace and remarkable thinness of the body 
(whence the name) at once arrest the attention. The great 
development of spines is also striking, there being two additional 
ones on the outer margin of the carpus of the cheliped and the 
same number on the anterior margin of the merus of the ambula- 
tory legs. 

Occurrence.—-The only specimen obtained was associated with 


P. maculata, and dredged among shells in 10-15 fathoms of water 
at the south end of the lake. 

3. General Remarks. 

The principal result of our extended knowledge of the Brachyura 
of the African lakes is to make still more clear the very special 
nature of the Crab-fauna of Tanganyika. As is found to be the 
case in so many different groups of animals, the forms occurring 
in Tanganyika are for the most part endemic, while those found 
in the other big lakes are often of wide distribution. There occur 
in Nyasa, as we have seen, three species of Potamonautes, one of 
which, it is true, is described from that lake only. Inthe Victoria 
Nyanza, we find Potamon (Parathelphusa) niloticus, a very widely 
distributed form, and P. (Geothelphusa) emini, also known from 
Abyssinia. From the Albert Edward Nyanza there comes also a 
species of Geothelphusa. Thus, with the single exception of 
P. (Potamonautes) orbitospinus, from Nyasa, the forms at present 
known from these big lakes are by no means confined to them. 

With Tanganyika it is quite otherwise. There are two species 
of Potamonautes, one of whichis not known elsewhere, but beyond 
this, three species of a unique and remarkable genus wholly re- 
stricted to this lake. From the other lakes we have then only 
representatives of the subgenera of Potamon—forms such as are 
widely distributed in the tropical fresh-waters of the Old World. 
From Tanganyika, while we have some representatives of these 
normal African types, we have a preponderance of forms perfectly 
distinct and occurring nowhere else. There is an indication, too, 
of that richness of the Tanganyika fauna which is so noticeable 
in some other animal groups. We know of five species of Crabs 
from Tanganyika, three from Nyasa, and only two from the 
Victoria Nyanza. 

It is necessary to add a few remarks on the affinities of the 
genus Platythelphusa. 'The species P. armata has been considered 
to exhibit a distinctly marine appearance. Milne-Edwards stated 
in the course of his original description that the Crab bore such a 
resemblance to certain. marine or brackish-water Grapside that 
we might relate it to that group, were it not for the development 
of the abdomen and the absence of metamorphosis. In his book 
on ‘The Tanganyika Problem,’ Moore* very rightly challenges 
the value of such a character as the latter to the systematist. If 
the absence of metamorphosis is the result of a particular habitat, 
as we have reason to suppose, we ought not to take it into account 
when we attempt to determine the aftinities of a newly-discovered 
specimen. At the same time it would seem as if the resemblance 
to the Grapside could be only a very superficial one, produced, 
perhaps, by the more or less quadrilateral shape of the carapace. 

* Op. cit. p. 236. 


It must be admitted that the considerable development of 
spines, which we find in all these species, is a feature more 
commonly found among marine Crabs than among those inhabiting 
fresh-water, and so would in itself convey a false impression on a 
casual inspection. But we need not go outside the subfamily of 
the Potamonine to find examples of Crabs which are just as 
“‘marine looking,” as far as the development of spines is concerned. 
This is true of a number of species of Parathelphusa, and when 
we pass to the other subfamilies of the Potamonide, we have 
Potamocarcinus, Valdivia, and Dilocarcinus, all extremely well 
armed with spines. As with all these genera and subgenera, 
there can be no doubt that Platythelphusa, despite its appearance, 
finds its nearest allies in this group of forms which are essentially 
and typically fresh-water in habitat. 

It is a matter of more difficulty to decide which of the allied 
forms are most closely related to Platythelphusa. The species now 
known as P. maculata has been stated to be a primitive form™*, 
on account of the little deflection of the front, the nature of the 
antenne, and the spine-bearing margins of the carapace. These 
features are equally characteristic of the other two species of the 
genus, and do seem to show less specialisation than the deflexed 
front and inflated gill-chambers of some of the semi-terrestrial 
species of the subgenus Potamon. It is then not with the 
latter, but near the genus Hydrothelphusa and the subgenus 

Parathelphusa that we ought probably to place this distinctive 
Tanganyikan genus. 

The problem of the origin of Lake Tanganyika, about which so 
much has been said and written, is of course intimately connected 
with the questions which have just been discussed. If the lake be 
the modified remains of part of an ancient ocean, we may expect 
its inhabitants to show both a primitive character and a marine 
aspect. These, Mr. Moore considers, are exhibited by all the 
members of his “ halolimnic” group, among which are reckoned 
the species of Platythelphusa. It has been shown, however, that 
‘this genus has no better claims to a marine origin than other 
representatives of the family, and at the same time, that while it 
is not so specialised as certain allied forms which have adopted a 
partially terrestrial mode of life, neither need it be considered as 
the most primitive in the group. Still there is little evidence 
that this form was ever anything but wholly aquatic, and it may 
have become modified by the truly oceanic conditions prevailing 
in Tanganyika, until it attained a superficial resemblance to 
marine types. 

In his report on the Macrurous Crustacea of the Expedition, 
Dr. Calman shows} that they are to be regarded rather as 
specialised than as primitive in character, and it may be asked 

* Proc. Zool. Soc. 1899, p. 702. 
+ Proc. Zool. Soc. 1906, p. 204. 


why the same is not true of the Brachyura. An explanation is 
really not far to seek. All the Macrura concerned are wholly 
aquatic types, while among the Brachyura we have to institute 
comparisons with forms which have partially accustomed them- 
selves to a terrestrial existence. Such a profound change in 
habits must produce an effect which, in comparison, would dwarf 
any modification brought about within the limits of a single 

There is gradually being accumulated a mass of information 
concerning the other animal groups inhabiting Tanganyika, and 
in nearly every case it is found that the forms are to a large 
extent endemic and, moreover, very distinct and highly modified. 
The explanation of this fact will be equally the explanation of 
the remarkable character and variety of the Tanganyika Crab- 
fauna, when compared with that of the other great African lakes. 
There seems little doubt that it is to be found in a growing 
divergence taking place in the lake during a prolonged period of 

Prate XVI. 

Fig. 1. Potamon (Potamonautes) orbitospinus (p. 259). Adult male, general view 
from above. Nat. size. 
Figs. 2-7. Series of figures of Platythelphusa and Potamon to show the frontal 
region and illustrate the relations between the front, descending process, 
and antenna (p. 267). 

Fig.2. Platythelphusa maculata. X 5. 
3. Platythelphusa coneulcata. X 5. 
4. Platythelphusa armata. X13. 
5. Potamon (Parathelphusa) niloticus. X 1s. 
6. Potamon (Potamonautes) platynotus. X 2. 
7. Potamon (Potamonautes) perlatus. X 2. 
Prate XVII. 
Fig. 1. Potamon (Potamonautes) platynotus (p. 264). Adult female, general view 
from above. Nat. size. 
2. Platythelphusa conculcata (p. 273). Male, general view from above. X 3. 
3. Potamon (Potamonautes) platynotus (p. 264). Larger chela of adult female, 
to show nature of dentation. X 1%. 
4. Platythelphusa conculcata (p. 273). Larger chela of male, to show nature 
of dentation. X 5. 
5. Platythelphusa maculata (p. 271). Chela of female, to show nature of 
dentation. xX 5. 
6, Platythelphusa maculata (p. 271). Abdominal region of a female, from 
below, to show the large size of the abdomen, and the degree to which 
it covers the sternum. X 3. 
Reference Letters. 
ant. 1. Antennule. ex.orb. External orbital spine. 
ant. 2. Antenna (second segment). Sub-orbital spine. Descending process of front. Sub-ocular tooth. 


5). On Two new Species of the African Genus Microchetus 
belonging to the Collection of Oligochzeta in the Museum 
of Christiania. By Frank H. Bepparp, M.A., F.R.S., 

Prosector to the Society. 
[Received February 1, 1907. | 
(Text-figures 85 & 86.) 

Dr. Robert Collett, the well-known chief official of the Christiania 
Museum, was so good as to entrust me, some little time since, 
with the collection of Oligocheta belonging to that Museum for 
study and description. In examining the collection I found 
three specimens, representing two species, of the Ethiopian genus 
Microchetus* which I believe to be new to science, and of which 
I beg to lay the following descriptions before the Society. 


I have the pleasure of dedicating this obviously new species to 
Dr. Collett. The material consists of but one specimen, which is 
entire, but considerably softened. It measures about 170 mm. 
in length by 7 mm. in breadth after the clitellum ; the anterior 
region of the body is wider. The colour is grey-brown, yellow 
on the clitellum. 

The sete have the usual paired arrangement found in this 
genus, and commence upon the second segment of the body. The 
sete are smaller upon the anterior segments and considerably 
larger in the clitellar region, where they are quite twice the 
length. The larger setz are ornamented. Some of the anterior 
segments consist of two rings. 

The clitellum (text-fig. 85) is very sharply marked off by its 
colour and by the greater thickness of the body-wall in this 
region. It commences with the xivth and ends with the xxivth 
segment. The position of the clitellum is not anomalous for the 
genus, and the specimen permits of no doubt upon the matter. 
The clitellum is perhaps best described as ‘“ saddle-shaped” ; but, 
as a matter of fact, the clitellar epithelium has also invaded the 
ventral surface. There is, however, a diminution in thickness 
indicated by an overhanging of the body-wall just ventralwards 
of the lateral sete. This arrangement is interfered with in the 
region of the genital papille by those structures, as will be seen 
by the accompanying figure (text-fig. 85). 

The nephridiopores are obvious and in front of the lateral pair 
of sete. 

The oviducal pores are very conspicuous upon the xivth segment. 
Each lies between and in the same line with the ventral and 
dorsal sete. 

* In addition to the species known up to the end of last century (see Michaelsen, 

Oligocheta in ‘Das Thierreich’), I. grisews has been subsequently described (see 
Michaelsen, MT. Mus. Hamburg; xix. 1904), 


The genital papille (see text-fig. 85) are large and conspicuous ; 
they occupy the xviith, xviiith, and xixth segments. They are 
quadrangular in shape, with rounded angles, and each is continued 
by a narrow ridge towards the middle ventral line. I traced the 
sperm-ducts in the interior of the body as far as the segments 
occupied by the genital papille, in connection with which I pre- 
sume that they open. But I am unable to fix more precisely the 
point of opening. 

Text-fig. 85. 

Microchetus colletti. 

T have not found it altogether easy, in view of the condition 
of the specimen and my unwillingness to injure it, as it is a 
type, to ascertain the position of certain of the internal organs. 
Assuming that the single pair of sperm-sacs is in the xth seg- 
ment, the strongly marked gizzard lies in segment vil. This 
segment is followed by two very thick septa, which thus separate 
segments Vil. /viii. and viii. /ix. The dorsal vessel is double in parts, 
and in segment ix. each half is much dilated, and a heart-like 
structure is formed precisely like that which I first described in 
Microchetus microchetus*, and which also occurs in other.species. 
The last pair of lateral contractile hearts les in segment xi. The 
calciferous glands are in segment 1x. 

The spermathece, which are, as is the case with other species 
of this genus, minute in size, open on to the boundary-line of 

* Trans. Zool. foc. xii. p. 63. 


segments x1./xil. and xii./xili. There are either three or four on 
each side of the nerve-cord in each segment scattered between 
the ventral region of the segment and the dorsal mid-line. 

Copulatory glands are rather numerous in this species. There 
are three pairs anteriorly, which le respectively in segments x., 
xi.,and xii. Each gland is double, as is commonly the case, being 
composed of two sausage-shaped glands uniting to form a common 
duct. They lie to the inside of the sperm-duct, which is almost 
or quite in contact with them as it passes back to the external 
pore. In addition to these three smaller pairs of glands there is 
a single much larger pair lying two or three segments behind the 
clitellum, and occupying a similar position in the body. ach of 
these glands i is also double; but each tubular half is longer and 
is coiled once upon itself. There are no conspicuous bundles of 
genital sete associated with any of these copulatory glands. 

The locality of the species is “ Zululand.” 


Two fragments contained in the same tube and from the same 
locality as the species just described cleariy belong to the same 
genus, but as plainly constitute another species of that genus 
hitherto undescribed. Both fragments include the entire head 
and body for a long way behind the clitellum. I do not give 
exact measurements; the diameter of the anterior part of the 
body is 12 mm. or so. The worms thus belong to a larger species 
than Microchetus colletti. 

One of the specimens (text-fig. 86) is more fully mature than 
the other, and there is a difference in the number of the genital 
papillee which is not attributable to immaturity. Iam thus able 
to give a better account of external characters. But as the worms 
are much softened my examination of the internal anatomy has 
led to less satisfactory results. Nevertheless, I have been able to 
make out certain anatomical facts which are of importance in the 
discrimination of species. 

The sete are so minute and difficult to see upon many segments 
that I have found it impossible to map the regions of the body 
by their aid only. Assuming, however, that the male pores are 
upon the border-line of segments xiv./xv.,a very usual position for 
them to occupy in this genus, and that the gizzard is in segment 
vil., which is also the case with other species, I arrive at the 
following determination of the position of the clitellum and of 
other organs. 

In the fully mature individual the clitellum, recognisable on 
the xivth segment, is completely developed on the xvth, and 
extends to the end of the xxviith. It is saddle-shaped, a bare 
ventral area being left. In addition to the clitellum, segments 
Xvi.-xxil. possess on either side a longitudinal band distinct from 
the clitellum, though of the same appearance in general, coupled 
with slight dissimilarity in colour, which is plainly the tubercula 


pubertatis. This is confirmed by the fact that in the second im- 
mature example without a clitellum the tubercula pubertatis were 
nevertheless quite plain, though apparently occupying a segment 

Text-fig. 86. 

Microchetus zuluensis. 

The copulatory pupille (text-fig. 86) with their sete are very 
obvious and very numerous in this species. In both specimens 
there are two series of these papillee, one anterior to the clitellum 
and one on some of the posterior segments of the clitellum. In 
the fully mature individual with the completely developed clitellum 
there is a single pair of these structures on segment x1., a papilla 
of the pair being situated a little to the dorsal side of each 
ventral pair of sete. These (the ventral) setee are also present on 
the xith segment. These anterior papille are considerably larger 
than those of the posterior series. The latter are in all five pairs 
situated on segments xxiii.-xxvii. inclusive, thus immediately 
continuing on the tubercula pubertatis. In the immature indi- 
vidual there were the same papille upon the xith segment ; but in 


addition to these there was a smaller pair upon the xiith segment, 
more ventrally placed, and in fact corresponding in position to 
the posterior series. These latter papille in the present specimen 
are fewer than in the adult, but they commence upon the same 
segment, 7. é. the xxilird. There are four upon the right side of 
the body and only two upon the left. 

The gizzard is distinctly contained in two segments, which are 
the vith and viith; the greater part of it, however, lies in seg- 
ment vil. The mesenteries separating segments vi./ix. are thick. 
The rather small calciferous glands are in segment 1x. ; the intes- 
tine begins suddenly in segment xi. The dorsal blood-vessel is 
dilated in segment ix. I imagine that this species is one of those 
which only possess one set of testes, funnels, &e. For the sperm- 
sacs and reservoirs consist of one pair in the ixth segment attached 
to the posterior wall of that segment and a pair in the xth 
attached to its anterior wall, 7. ¢. the same septum as that which 
bears the sperm-sacs. The two sacs of each side of the body seem 
to communicate, and I take the anterior pair to be the sperm-sacs 
and the posterior pair to contain the funnels. 

T was quite unable to find any spermathece. 1 am unwilling, 
however, to assert that these organs are absent. If they happened 
to contain no sperm their minute size and the softened condition 
of the worm would render it at least very difficult to detect them. 
The only other species, as it appears, in which no spermathecz 
have been detected is Dr. Michaelsen’srecently described I. grisews*. 
But as this latter species is holandrous it cannot be confused with 
M. zuluensis, than which it is also a good deal smaller, 

March 19, 1907. 

Dr. Henry Woopwarp, F.R.S., Vice-President, in the Chair. 

The Secretary read the following report on the additions that 
had been made to the Society’s Menagerie in February 1907 :— 

The registered additions to the Society’s Menagerie during the 
month of February were 76 in number. Of these 27 were 
acquired by presentation and 10 by purchase, 36 were received 
on deposit, 1 by exchange, and 2 were born in the Gardens. 
The total number of departures during the same period, by death 
and removals, was 188. 

Among the additions special attention may be directed to :— 

A Long-tailed Goral (Wemorhedus caudatus), from Korea, new 
to the Collection, presented by Mr. C. F.G. Bilbrough on Feb. 5th. 

A Harpy Hagle (Thrasaétus harpyia) from South America, 
purchased on Feb. 8th. 

Myr. Herbert F. Standing read a paper, illustrated by lantern- 

* Loc. cit. (on p. 277). 


slides and large series of photographs and specimens, on recently 
discovered subfossil Prosimiz from Madagascar, in which he dis- 
cussed their affinities with extant Lemurs and with the higher 
Primates. The remains were obtained in the muddy bed of a swamp 
formed by the blocking-up of the river Mazy by a lava-flow, at from 
a few inches to 3 or 4 feet below the surface. They consisted of 
a large number of skulls and limb-bones of Lemurs and Lemur- 
like animals. This great amount of material enabled the author 
to corroborate the view, previously put forward by Dr. Forsyth 
Major, that the extinct Lemurs of Madagascar were, in many 
respects, intermediate between existing Lemurs and Monkeys, 
and to express his belief that the New World Monkeys and the 
Lemuride, as well as the Malagasy Indrisine, had a common origin. 
He also stated his opinion that, in view of the recent additions to 
our knowledge of the Prosimiz and of what the present collection 
revealed with regard to their close relationship to the Apes, it 
was not possible to separate the Primates, as hitherto, into the 
two suborders Lemuroidea and Anthropoidea. 
This paper will be printed entire in the ‘Transactions,’ 

The following papers were read :— 

1. Descriptions of some New Species of Animal Parasites. 

By L. W. Sampon, M.D., F.Z.S.* 

[Received March 19, 1907. } 


Abstr. P. Z. 8. 1907, p. 15 (March 26). 

Habitat. Small intestine of Pedetes cafer. 

Only females found, 12-15 mm. long and about 1 mm. broad. 
Characterised by the presence of a conical ovipositor 2-3 mm. 
long, placed ventrally on the anterior third of the body, 2- 
3°5 mm. from the cephalic extremity, and by a spirally twisted 
tail, 2-3 mm. long, terminating in a fine point. Body semitrans- 
parent. Head tapering anteriorly. Mouth trilabiate ; cesophagus 
long and terminating in a spherical bulb. Anus open ventrally 
at 3-4 mm. from tail-end. Eggs smooth, oblong, asymmetrical, 
and measuring 60-65 p by 28-32 p. 


Abstr. P. Z. 8. 1907, p. 16 (March 26). 

Habitat. Connective tissue of Man. 
Long, flat, unsegmented body, 15 em. long and 1:5 mm. broad, 

* [The complete account of the new species described in this communication 
appears here; but since the names and preliminary diagnoses were published in the 
« Abstract,’ the species are distinguished by the name being underlined.—Ep1ror. | 


with numerous irregular transverse folds and a distinct longitu- 
dinal groove on ventral surface. Anterior extremity 2-5 mm. 
broad; head completely invaginated. Posterior extremity 2 mm. 
broad, with shallow median slit. Extracted from an abscess on 
the thigh of a Masai in British Central Africa. 


Abstr. P. Z. 8. 1907, p. 16 (March 26). 

Habitat. Blood-vessels of Man. 

In the Congo Free State, in other parts of Africa, and in the 
West Indies there is a form of Bilharziasis clinically and patho- 
logically similar to the Asiatic form caused by Schistosomum 
japonicum, and unlike the classic Kast African form due to 
S. hematobium. The eggs of the species which causes this peculiar 
form are never found in the urine, but seem to be eliminated 
through the intestine only. They differ from those of S. hema- 
tobiuwm in having a broad lateral spine totally different in size, 
shape, and position from the small, straight, terminal spe which 
characterises the ova of S. hematobiwm. Hitherto, the laterally 
spined ova, usually observed in Egypt in cases of mixed infection, 
have been looked upon as having been distorted while passing 
through the rectal mucosa. Sir Patrick Manson suggested several 
years ago, that the laterally spined ova found in the feces of 
patients, and never in the urine, might represent a new species. 
In appreciation of this, one of his many genial intuitions, the new 
species 1s dedicated to him. 

2. Descriptions of five New Species of Heemogregarines from 
Snakes. By L. W. Samson, M.D., F.Z.8S., and C. G. 
SeLicmMann,. M.D., F.Z.8.* 

[ Received March 19, 1907. | 


"Abstr. P. Z. 8. 1907, p. 16 (March 26). 

Habitat. Erythrocytes of Indian Python, Python molurus L. 
Club-shaped, 14-16 p long. Anterior extremity rounded, 3-15 
broad. Posterior extremity attenuated and recurved. Cytoplasm 
more or less granular. Nucleus median or nearer posterior ex- 
tremity, large, oval, and with coarse, deeply staining chromatin 
granules. Parasite lies parallel or obliquely to long axis of host- 
cell, of which it occupies about two-thirds, without causing much 

alteration beyond displacement of nucleus. 

* [The complete account of the new species described in this communication 
appears here; but since the names and preliminary diagnoses were published in the 
* Abstract,’ the species are distinguished by the name being underlined.—Eprrokr. | 



Abstr. P. Z. 8. 1907, p. 17 (March 26). 

Habitat. Erythrocytes of Diamond Snake, Python  spilotes 

Club-shaped. Some forms more slender, 14-15 long and 2 yu 
broad, with both extremities rounded and differing only slightly 
in thickness. Other forms more pay somewhat similar to those 
of Python molurus, but larger, 22 uw by 4. Nucleus median 
and very large, 9p by 4p. Host-cell sometimes slightly distorted, 
nucleus pushed to the periphery. 


Abstr. P. Z. S. 1907, p. 17 (March 26). 

Habitat. Erythrocytes of Hoary Snake, Pseudaspis cana L. 

Crescentic, bean-shaped, and discoidal forms occur. The slender 
crescentic forms have a long, oval, and more or less central nucleus. 
The bean-shaped forms measure 10-12 y in length by 5-6 « in 
width; they have a wide central nucleus, and their cytoplasm is 
literally crammed with rounded, highly refractive granules. Host- 
cell unaltered beyond occasional displacement of nucleus. 


Abstr, P. Z. 8. 1907, p. 17 (March 26). 

Habitat. Erythrocytes of Testaceous Snake, Zamenis flagelli- 
Forms L. 

Oval or bean-shaped cyst 12-13 » long by 5-6 w broad, en- 
closing club-shaped parasite doubled up in the form of a letter U 
with both branches of equal length and closely applied. Nucleus 
median and situated near bend at one pole of cyst. Chromatin 
arranged in transverse parallel lines or in concentric circles. 
A characteristic feature is the almost constant presence of two 
large chromatoid granules usually placed one on each side of 
nucleus. Host-cell unaltered, nucleus slightly displaced. 


Abstr. PB. Z. 5. 1907, p. 17 (March: 26): 

Habitat. Erythrocytes and leucocytes of Couper’s Snake, 
Coluber corais var. couperi Holbr. 

1. Slender, elongate, cylindrical forms 14 p long by 1:5 p broad, 
sometimes presenting a refringent granule or vacuole at each 
extremity. Host-cell apparently unaltered or only slightly 
Pear trophied. 

Large bean-shaped forms 12-13 pw long by 4-5 pw broad, with 
Res more or less granular and occasionally vacuolated. 
Nucleus small, round, median! with fine chromatin grains. The 
host-cells containing ‘this form measure about four times the 
normal size, and are entirely dehzemoglobinized and greatly atten- 
uated. Their nucleus is hypertrophied. Sometimes two or even 
three parasites may be found in the same host-cell. 


3, The Rudd Exploration of South Africa.—VII. List of 
Mammals obtained by Mr. Grant at Coguno, Inham- 
bane. By OLp¥FieLp THomas, F.R.S., F.Z.8., and R. ©. 
Wrovuenton, F.Z.8. 

[Received February 21, 1907. ] 

No part of South Africa was more inadequately represented in 
our National Museum than the Portuguese territories north-east 
of Delagoa Bay, and we are glad to say that the fine collection 
from Inhambane of which we now give an account does much 
towards filling up the lacune. Mr. Grant found a good collecting 
place at Coguno, about 75 miles to the south-west of the town of 
Inhambane, and obtained there the whole of the present series, 
which numbers. 212 specimens, belonging to 59 species. 

Of these we have described 6 as new, several of them being 
particularly striking forms, notably a Galago and a Petrodromus. 

And besides the novelties, specimens that are of very special 
value are the topotypes of such of Peters’s species as were described 
from Inhambane, these being the first of a series of Petersian 
topotypes which we hope Mr. Grant may in time be able to com- 
plete, since all S.E. African Mammalogy seems to circle round the 
species obtained during the epoch-making voyage of the famous 
German zoologist, and but little satisfactory work can be done until 
good modern specimens of all his species are available for study. 

As before the whole of this valuable collection is presented to 
the National Museum by Mr. C. D. Rudd. 

Mr. Grant’s notes on the Inhambane District are as follows :— 

“Tnland from Inhambane the country is composed of more or 
less undulating, sandy flats, densely bushed and timbered ; often 
for many miles the only open spaces are the clearings made by the 
natives for their kraals and cultivated lands. 

“Some of the river-valleys are more open, consisting of patches 
and stretches of forest alternating with open plains, generally 
thickly covered with palmetto. 

“The rivers, excepting the Intanime, are merely huge, dense 
reed-beds, through which the actual watercourse is often traceable 
only with difficulty. 

“The natives are very numerous and are mostly Mchopee, with 
a small sprinkling of Machangaan, and all are great hunters and 

trappers, and gave considerable help throughout the trip, either 
by giving information as regards species or in bringing in 
‘My head camp lay close to the Commando of Coguno, which 
is about 130 kilometres by the Panda Road from Inhambane ; : 
thence short trips and excursions were made into the surrounding 
country as necessity demanded. 

“Owing to the denseness of the bush and the comparative 

flatness of the veldt local guides were always necessary. 

Proc. Zoou. Soc.—1907, No. XX. 20 


‘“¢So far as I could learn, the District has never been surveyed, 
so that the altitude of Coguno, or other places, is unknown, 
but probably no part exceeds 1000 ft. The actual position of 
rivers and localities can only be roughly estimated. 

“Throughout my stay the climate was delightful ; the average 
temperature being around 80° in the shade, occasionally reaching 
88°; it being the dry season, there was little rain. 

“The following species were not obtained but exist at Coguno, 
and the native names may be of interest. They are: 

WING OS ORGCUIS peer eres ee: Lengo. 
Thryonomys swinderenianus ... Shleti.,.—C. H. B. G. 

1. Papro porcArius Bodd. 
Flat skin. 

“‘ Native name, ‘ Infeni.’ 

“ This species was not seen in the neighbourhood of Coguno 
Camp, but is said to be common to the North and West. The 
flat skin sent was obtained from a native who killed it some 
20 miles to the northward of my camp.”—C. H. B. G. 


Mr. Pocock informs us that he considers Peters’s erythrarchus, 
of which these are topotypes, should be ranked as a subspecies of 
pygerythrus F. Cuvy. 

“ Native name, ‘ Makaku’ or ‘ Ihow.’ 

“¢Gommon and inhabiting the denser forests where it lives on 
the wild fruit and berries. Never observed in large troops ; single 
old males were sometimes seen, but were usually unapproachable. 
They are at all times wary and extremely difficult to secure. 
Cercopithecus albigularis exists in the district, but is very rare 
and local and even the natives were unable to obtain me speci- 
mens; it is known to them as ‘ Glanglanu ’.”C. H. B. G. 

3. GALAGO GRANTI, sp. n. 
3. 1617, 1618, 1619, 1660, 1662. Q. 1517, 1653, 1663. 

A member of the moholt group, with an unusually bushy black- 
tipped tail. 

Fur fine and soft, about 15 mm. in length on the back. General 
colour above drab-brown, rather darker than Ridgway’s “ drab.” 
Under surface cream-buff, the basal three-fifths of the hairs slaty. 
Light nose-line and black orbital patches as in G. moholi. Outer 
side of fore limbs light drab, lightening to white on the hands ; 
inner side like belly. Hind limbs as usual dull cream-buff 
throughout, rather duller than on the belly. Tail long, unusually 
bushy, the hairs attaining a length of 20-25 mm., those of true 
moholi but little more than half this length ; in colour the basal 
three-fifths of the tail is drab-brown like the body, gradually 
darkening terminally to blackish brown, 


Skull readily distinguishable from that of G. moholi by its much 
longer muzzle, the palate-length being over 17 mm., as com- 
pared with 15 in the type of moholi, and all the other muzzle 
measurements in proportion. The bulle also are lower and 
less abruptly swollen anteriorly, and the canines appear to be 
rather heavier. 

Dimensions of the type :— 

Head and body 158 mm.; tail 237 ; hind foot 63; ear 43. 

Skull—greatest length 45 mm.; basal length 35; zygomatic 
breadth 28; mastoid breadth 23°5; tip of nasals to back of 
orbits 25; palate-length 18; front of canine to back of m® 15-6. 

Type. Adult male. Original number 1662. 
Collected 26 August, 1906. 

This Galago differs from G'. moholi by its long bushy black- 
tipped tail and by its markedly longer muzzle. Peters’s G. mos- 
sambicus, the skull of which is figured by him under the name of 
senegalensis, has a short muzzle as in the true moholi. 

With regard to G. conspicillatus I. Geoft., Prof. Trouessart 
kindly informs us that the tail of the type is only 20 mm. in total 
breadth, therefore exactly as in moholi, not as in the Inhambane 

We have had great pleasure in naming this interesting little 
‘‘ Bush-baby ” after its captor, Mr. C. H. B. Grant, who has 
collected the whole of the immense number of mammals recorded in 
the present series of papers, and has thus ably utilised the splendid 
opportunity afforded him by the generosity of Mr. Rudd. 

“ Native name, ‘Suwanjati.’ 

‘¢Common and inhabiting the forests. It is strictly nocturnal, 
sleeping during the day in the hollow trees, where it may generally 
be taken in small family parties. ‘This species like many others 
is eaten by the natives. 

“ G. crassicaudatus was several times heard calling in these 
parts at night, and I saw one skin worn by a boy; none, however, 
- could be secured by myself or the natives, although I offered good 

rewards fora specimen. It is known to the natives as ‘ Gerile ’.” — 

(Op Jal, 16) (Cr 


$. 1577, 1578, 1621,1656. 9. 1579, 1616, 1665, 1671, 1672, 

In this series there would seem to be two forms distinguished 
by size. The larger corresponding closely to, if not identical with, 
the specimens from Klein Letaba provisionally identified by 
Thomas & Schwann (P. Z. 8. 1906, p. 577) as migrita. The 
presence in one place of two forms differing in size seems to be a 
common occurrence in this group, as for instance, planirostris and 
viridis of Peters from the Zambesi Valley, and nigrita herero 
and damarensis of Thomas from 8. W. Africa. We are of opinion 
that it is safer to include all these specimens provisionally under 

nigrita until the group can be worked out in its entirety. 


“‘ Native name, ‘ Mongavilane.’ 

“Quite the commonest of the Bats at Coguno. Generally 
appears long before dark, flying fairly high, and can be easily 
secured with a shot-gun. Numbers 1621 and 1673 appeared to 
me different from the others, from which they were easily 
distinguishable on the wing.”—C. H. B. G. 

Go UDO, Gaile 

This is the most southern locality from which this small Scoto- 
philus has been recorded, the next being Marungu, whence 
Dr. Noack described his S. minimus. 


¢. 1580, 1603. ©. 1620. 

Additional specimens of the rare and remarkable Butterfly- 
bats of the genus Glawconycteris are extremely welcome. The 
present examples agree in size and the colour of their heads with 
G. papilio, and equally differ from the white-headed and white- 
bellied G. variegatus Tomes, of Damara-land. 

“‘ Native name, ‘ Mongavilane.’ 

“ Apparently uncommon, as the three specimens taken were 
the only ones observed. They appear about the same time as 
Scotophilus, but have a more butterfly-like flight, which easily 
distinguishes the species at a good distance.” —C. H. B. G. 

©. 1668. 

Quite like the specimens from Legogot mentioned in the last 
paper on the Rudd Collections (Thomas & Schwann, P. Z. Sh 
1906, p. 780). 

“ Native name, ‘ Mongavilane.’ 

“These little Bats were fairly common, but, owing to their 
small size and rather late appearance, a big series was not 

obtainable.”—C. H. B. G. 


B Ws, IGS7, WIM, De Webs) 

“Caught in run in thick bush.” 

These valuable specimens are practically topotypes of Peters’s 
species, which was described from the neighbourhood of Inhambane 

Their bellies are of a peculiar coppery brown, and the lighter 
basal portion of their dorsal hairs is also strongly tinged with the 
same colour. In A. chrysillus Thos. & Schw., from Delagoa Bay 
the same parts are white or yellowish white. The hairs of the 
back are about 5 mm. long in 4. obtusirosiris, 6-5 mm. in 

A. chrysillus. 
* Ann. & Mags. N. Hi. (7) xv. p. 77, 1905. 


“* Native name, ‘ Tukunve.’ 

“ Fairly common and forming runs just below the surface in 
the sandy ground, in the forest and bush. Owing to the peculiar 
nature and smallness of the runs it was found impossible to trap 
it and it could only be taken when seen working.”—C. H. B. G. 


6. 1521, 1523, 1536 1539, 1557. 9. 1500, 1515, 1522, 1524, 
1537, 1538, 1552, 1553, 1554, 1555, 1556. 

Allied to P. sultan* Thos., but skull shorter and the tail more 
hairy and more finely scaled. 

General characters as in P. sultan, the peculiar round-headed 
bristles underneath the tail quite as in that species. Colour 
sunilar throughout, except that the dull russet-brown dorsal area 
is broader and more diffused, extending nearly across the back, 
instead of forming a comparatively narrow line down the spine 
sharply separated from the grey of the shoulders and flanks. 
Face-markings, limbs, and under surface all quite as in sultan. 

Tail rather longer than in suléan, and the scaling finer, 13 rings 
of scales to the centimetre instead of 10 as in sultan; upper 
surface well haired throughout, the scales nearly or quite 
hidden—(in P. sultan the upper side of the tail is practically 
naked and the large scales are clearly visible). In colour the 
upper surface is deep black throughout, and the lower dull buffy, 
not sharply contrasted; extreme base below pale flesh-colour. 
Long knob-headed bristles black throughout. 

Skull markedly shorter than in P. swltan, but the shortening is 
chiefly in the muzzle, the brain-case being of about the same size. 
Hinder edge of nasals level with the front of the anteorbital rim, 
and falling about one millimetre short of the frontal processes of 
the maxille. In P. sultan the nasals are of about the same 
length, but end further forward. P* (the fourth tooth from the 
back) comparatively small and narrow. 

Dimensions of the type (measured in the flesh) :— 

Head and body 192 mm.; tail 181; hind foot 57; ear 36. 

Skull—greatest length 53 mm.; basal length 47; greatest 
breadth 29°5; length of nasals 20; interorbital breadth 9; 
breadth of brain-case 20; length of upper tooth-series 27; front 
of p’ to back of m* 11°5. 

Type. Old male. B.M. no. Original number 1557. 
Collected 5 July, 1906. 

The occurrence of this handsome Petrodromus in Inhambane 
was already recorded by W. L. Sclater?, who had received a 
specimen collected there by Mr. H. F. Francis. With only a 
single specimen, however, he did not think himself justified in 
distinguishing it from P. sultan, and it is therefore placed in the 
‘Mammals of South Africa’ under the latter name. 

* P.Z.S. 1897, p. 435. Originally published as P. suJtani by a printer's error, 

but the mistake was corrected P. Z.S. 1897, p. 928. 
+ Mainm. S. Afr. 11. p. 155, 1901. 


In Herr O. Neumann’s revision of the genus* the primary 
division of the species is made by the hairiness or nakedness of 
the upper side of the tail, but the occurrence of this species, 
obviously a close ally of P. sultan but with a hairy tail, shows that 
this character is of but little importance. We should rather 
divide the species, as in Thomas’s original paper, by the structure 
of the caudal bristles, in which respect P. schwanni agrees with 
P. sultan alone of all the described forms. 

No Petrodromi of this type have been as yet recorded between 
Inhambane and the mainland opposite Zanzibar, a distance of 
some 1200 miles. 

Weare glad of the opportunity of linking with this fine species 
the name of Mr. Harold Schwann, who has hitherto been so 
closely connected with the working out of the Rudd Collection. 

‘“‘ Native name, ‘ Nyakole.’ 

“Very common and inhabiting only the dense thickets, where 
it has regular runs, in which it is easily trapped. It has all the 
actions of the Macroscelidide, carrying the tail almost perpen- 
dicular when running. Exclusively diurnal and insectivorous.”— 

CoH BG: 

10. Crocipura sytv1A Thos. & Schw. 

6. Ldai3: 

“ Native name, ‘ Nongi’ (without distinguishing species). 

‘* Not common and confined to the reed-beds and swamps along 
the river-valleys, especially the Inyasuni.’—C. H. B.G. 

Ge V5e2 1574: 

12. Fevis ocrEATA Gmel. 
2. 1636. 

We hesitate to identify this single specimen with any particular 
subspecies until the arrival of further specimens from Portuguese 
Kast Africa. 

““ Native name, ‘ Igoye’ or ‘Simange.’ 

‘“‘ Apparently very uncommon, as the specimen obtained was 
the only one taken or observed.”—C. H. B. G. 

OP One 

“‘ Native name, ‘ Fungwi.’ 

“‘ According to the natives common, although I did not observe 
more than the one specimen, except a couple of skins in the 
possession of natives. Apparently inhabits the thickest of 
forests.” —C. H. B. G. 

* Zool. Jahrb., Syst. xiii. p. 541, 1900. 


14. GENETTA sp. 
3g. 1566. @. 1562. 

Allied to G. letabe Thos. & Schw. 

‘“‘ Native name, ‘Simba.’ 

** Not very common and found everywhere, especially near 
native habitations. This species, with all the cats, is eaten by 
the natives.”—C. H. B. G. 

15. Muncos GALERA Erxleb. 

3. 1645. 

“* Native name, ‘ Shikoko.’ 

“« Apparently common and generally living in the reed-beds and 
swamps; the specimen sent, however, was caught some little 
distance from water and near a Kaftir kraal, where it was 
probably on the prowl after a stray chicken. Nocturnal only.”— 

C. H. B.G. 

16. Muneos (IcHNEUMIA) GRANDIS Thos. 
3. 1535. 

Whether this animal, the external characters of which are quite 
as in southern examples of MW. albicauda, should be treated as one 
of the subspecies of that form, or whether several of these should 
be raised to specific rank, is a question which cannot be settled 
without further material. Immature specimens would be of 
particular value as the characters rest mainly on the structure 
of the teeth, which get worn down in adult life. 

‘‘ Native name, ‘Sanganye.’ 

“By no means common and generally found near habitations, 
which they visit for the chickens, &c.”—C. H. B.G. 

17. MunGos CAFER Gm. 

“Native name, ‘ Shlaushlwa.’ 

_ The flat skin sent was the only one seen during the trip. It 
is evidently rare, as many of the natives did not know its name 

and some even had never before seen one. The boy from whom 

I took it said he caught it in the reedy bed of the Inyamatanda 

River.”—C. H. B. G. 


3. 1558, 1628, 1654. 

“Shot in thick bush.”—C. H. B. G. 

Much greyer than the true fasciatus. 

Size rather larger than in Zululand examples of fasciatus, 
approaching that of the Ruwenzori form, C. f. macrurus Thos., 
but the tail not lengthened. General colour conspicuously greyer 
than in fasciatus, the nape, fore-back and flanks of a clear 
cinereous grey, entirely without fulvous suffusion. The stripes 
normal in number and position, but the light ones white along 


their anterior edge, only becoming a little fulvous posteriorly ; 
in fasciatus each light stripe 1s fulvous anter iorly becoming darker 
fulvous poster iorly. Head finely grizzled grey, blacker on the top 
of the muzzle. Ears grey, without fulvous suffusion. Limbs 
grizzled grey, dar kening terminally on the hands and feet to 

Skull of normal proportions, but the teeth unusually large. 

Dimensions of the type (measured in the flesh) :— 

Head and body 364 mm.; tail 236; hind foot 73; ear 24. 

Skull—condylo-basal length 73 mm.; basal length 68; zygo- 
matic breadth 39; palate- length 39; greatest diameter of p 74. 

Type. Adult male. B.M. no. 6. 11.8.51. Or igimal number 
1628. Collected 13 August, 1906. 

This beautiful grey form of the Striped Mungoose may be 
readily distinguished by the reduction of the fulvous suffusion of 
the fur, this being only present along the posterior bor der of each 
dorsal light band, while in the orher forms some trace of it 
oceurs all over the body; the clear grey nape and shoulders of 
C. f. senescens are especially noticeable. 

The young specimen no. 1558 is more like the typical form. 

‘“‘ Native name, ‘ Gale.’ 

‘“* Apparently fairly common, but dithcult to secure owing 
partly to its wariness and partly to its often inhabiting the 
denser parts of the bush. The specimens obtained were shot 
during native hunts and were taken from troops of perhaps eight 
individuals. It is distinctly gregarious and diurnal in habits, 
living principally on coleopterous insects and the eggs and young 
of ground-breeding birds.” —C. H. B. G. 

‘“‘Native name, ‘ Sididi.’ ”—C. H. B. G. 

20. Icronyx CAPENSIS Kaup. 
1534, 1655. 9. 1575 
‘“‘ Native name, ‘Shingemani.’ 
“Fairly common everywhere, especially near native kraals, 

where it makes itself a considerable nuisance by stealing chickens. 
Nocturnal only.”—C. H. B. G. 


3. 1596, 1600, 1608 (yg.), 1609, 1610. 9. 1547, 1560, 1561 
1611, 161 2. 

A Funisciurus intermediate in size between cepapi and palliatus, 
with the colour pattern of the former and the bright colouring of 
the latter. 

Size rather smaller than in /’. palliatus. 

Fur of back soft, 12 mm. long; that of tail 30-40 mm. 

General colour above erey-br own freely grizzled with very pale 
buff or yellow; the individual hairs black or dark brown, each 


with two rings (one near base and one subterminal) yellowish 
white ; the belly orange-red, the hairs unicoloured to their bases. 
Head igen the forehead backwards coloured the same as the back ; 
face below the eyes, chin and throat orange like the belly. Outer 
side of limbs coloured like the back, but a strong rufous tinge on 
thighs; inner side orange like the belly. Tail coloured like the 
back but with a strong rufous tinge at the sides and below; the 
individual hairs with two pale rings, each 5 mm. wide, separating 
three black rings of same width, with a long (10-12 mm.) ferru- 
ginous tip. 

Skull as in palliatus but decidedly smaller. 

The following are measurements of the type (which scarcely 
differ from those of others of the series). For convenience of 
comparison corresponding measurements of a normal adult speci- 
men of F, palliatus ornatus Gray from Zululand are added between 

Head and body 197 (200) mm.; tail (cire.) 195 (218); hind foot 
41 (45); car 20 (18). 

Skull—greatest length 48 (52) mm.; basilar length 37 (40); 
zygomatic breadth 28 (30); interorbital breadth 15°5 ml 5); nasals 
13°7 (15°5); upper molar series 8°7 (9°5); bulle 10°5 (11). 

Type. Adult female. B.M. no. Original number 
1560. Collected 5 July, 1906. 

Through the courtesy of Dr. Péringuey we have been able to 
examine a Squirrel from St. Lucia Bay, Zululand (South African 
Museum, no. 4361), assigned by Mr. W. Sclater * to frerei Gray, 
under the impression that the type of the latter was obtained by 
Sir Bartle Frere in Natal. But the typical specimen was received 
by the British Museum in 1873, about which date Sir B. Frere 
veturned to England from Zanzibar, and he did not go to South 
Africa till some years later. The type locality “‘ Zanzibar” given 
by Gray for his “ Macroxus anumulatus frere” is therefore 
undoubtedly correct. The Zululand specimen, though paler in 
colour, agrees in all essential characters with the present species, 
to which we have no hesitation in assigning it. The range 
of sponsus is therefore from Inhambane to Zululand, where it 
coexists with F. palliatus ornatus Gray, and this fact and its 
smaller size ee justify us in classing it as a distinct species. 

‘““ Native name, ‘ Shintsi.’ 

““ Extremely common and found everywhere in the forests and 
thickets, especially near native clearings; as many as half a 
dozen can be seen at one time running about in the trees. The 
alarm call is a bird-like chatter. It is taken in vast numbers by 
the natives, with whom it is an especial delicacy.” —C. H. B. G. 

As we have had occasion to refer to the true locality of frerei, 
we further take this opportunity of recording that, allowing for 
the fact that it is a young individual, we find on comparison that 
the type of frerci agrees quite closely with the specimens assigned 

* Mamma. S. Afr. ii. p. 7 (footnote), 1901. 


by Herr Neumann to his /. palliatus suahelicus from the same 
region, and we have no doubt as to their identity; swahelicus 
Neum. must therefore give way to the older frerei Gray. 


3. 1484, 1488, 1491, 1499, 1531,1532. 9. 1483, 1487, 1489, 
1494, 1514, 1533. 

These specimens, though less pale than typical bechuane from 
the Bechuana Desert, are very close to that form. Their receipt 
from Inhambane is interesting from the point of view of the 
distribution of the species. The type of lobengule was described 
from Matabililand and local races were subsequently named from 
Kuruman, Matopo, and Salisbury, all places which may bedescribed 
as on the left bank of the Limpopo. In the collection from the 
Zoutpansberg District on the right bank, though more than one 
species of 7atera were found, there was no specimen referable to 
lobengule. More recently, however, in a small coljection from 
Pietersberg District on the right bank of the Limpopo but much 
lower down than Zoutpansberg, some specimens scarcely distin- 
guishable from the present series were found. Broadly, however, 
the area between the Limpopo and Zambesi Rivers would seem to 
be the home of the species, which takes quite well-marked local 
forms in various parts of its habitat. 

‘“‘ Native name, ‘ Singaan.’ 

“ Quite the commonest rat in the district and found everywhere, 
especially in and around the native clearings.”—C. H. B. G. 


3. 1583, 1584, 1586. 2. 1585, 1587, 1588. 

These specimens are all young but agree closely with O. irroratus 
cupreus from Zoutpansberg. 

‘“¢ Native name, ‘ Woti.’ 

“Not common and only taken in the reed-bed and swamps 
along the river.”—C. H. B. G. 


do. 1498, 1519, 1614, 1625, 1626, 1630, 1634,1647. 2. 1486, 
1607, 1615, 1623, 1633, 1638. 

“ Native name, ‘ Maklangane.’ 

“Tt is here far commoner than in any locality where I have 
previously taken it. Found everywhere, both in the forest and 
bush and the native lands, where it has regular tracks and runs, 
sunilar toits congener at the Klein Letaba. Exclusively diurnal.” 

—C€. H. B. G. 


3. 1495, 1505, 1506, 1508, 1509, 1597, 1606, 1629, 1639. 
Q@. 1479, 1481, 1482, 1490, 1501. 

These specimens are practically topotypes of S. fwscus Peters, 


described from Inhambane, but as they are indistinguishable from 
S. campestris, it seems probable that fuscus was based on a dark 
coloured specimen and represents no more than an individual 

“ Native name, ‘Sikwikle.’ 

‘Very common and frequenting the native lands and the 
adjacent bush.”—C. H. B. G. 

26. Mus ratrus L. 

6. 1520, 1526, 1527, 1551, 1581, 1652, 1657, 1658. 9. 1528. 

‘¢ Native name, ‘ Tikonso.’ 

‘¢ Very common and confined to the kraals and houses, never 
apparently visiting even the adjacent cultivated land. Several 
other examples besides the specimens sent were seen and all were 
very smallin size. Apparently exclusively nocturnal.”—C. H.B.G. 

27. Mus coucHa Smith. 
6. 1497, 1502, 1511, 1512, 1605, 1624, 1666. 9. 1480, 
1504, 1648. 

‘¢ Native name, ‘Supwisne.’ 
‘¢Common and habits similar to those of members of the species 
in other parts of 8. Africa.”—C, H. B. G. 

28. Mus cHRYSOPHILUS de Wint. 
ge Lo0d, Lol. tol3) 1529, 15485 L60l) Os 1530) 1540) 
Toa AG a a0: 

‘Native name, ‘ Sinse.’ 
““Common ; habits similar to those of members of the species 
in other parts of 8. Afvica.”—C. H. B. G. 


6. 1604, 1643. ©. 1644. 

‘¢ Native name, ‘ Shikoloveta 

“« According to the natives this species is fairly plentiful, although 
they were unable to get me specimens ; the three secured being the 
only examples I saw. It is arboreal in habits, ferming, according 
to native report, nests similar to Dendromus or Th. dolichurus.” 

—C. H. B.G. 


3. 1490. 

“‘ Native name, ‘ Senbendenyumbe " 
‘“‘ Apparently rare, as the specimen sent was the only one taken 
or observed.”—C. H. B. G. 

6. 1632, 1641, 1651. 2. 1642, 1650, 1659. 
Size and general characters as in gambianus and g. viator; the 


body-colour above much as in viator. ‘The sharp line of demareca- 
tion between the colour of the back and the white of the belly, 
which is strongly pronounced in typical gambianus, and plainly 
though less markedly so in viator, completely absent; the cheeks 
and flanks lighter than in viator, the lower part of the former white 
like the belly. Otherwise the colour-pattern quite as in viator. 

Skull in size and other essential characters as in viator ; but the 
upper molar series rather stronger, the anteorbital foramina 
slightly broader, and the rudimentary postorbital processes more 

Dimensions of the type :— 

Head and body 358 mm.; tail 438; hind foot 71; ear 40. 

Skull—greatest length 75 mm.; basilar length 65; zygomatic 
breadth 36; nasals 3211; interorbital breadth 11°8; palatal 
foramina 8°6; length of upper molar series 11. 

Type. Adult male. B.M. no. Original number 
1632. Collected 14 Aug. 1906. 

These specimens are from the 8.E. limit of the range of the 
species, and it is consequently not surprising to find such small 
differences as those recorded above between this and the Nyasa 
form, from which it is separated by nearly two degrees of latitude. 

‘“‘ Native name, ‘Sigwinye.’ 

‘“‘ Fairly common and generally found in thickets and densely 
wooded places, where it forms burrows of 2-6 holes, almost 
indistinguishable from those of Pedetes cafer. It is exclusively 
nocturnal in habits, and usually is only to be obtained during 
the dark phase of the moon; on moonlight nights it seldom comes 
out, owing, according to the natives, to its great fear of the larger 
owls. During the dark nights it lays in a store of food to tide it 
over the moonlight nights, until it can again venture out. I have 
observed that it also loosely fills the entrance of the burrow with 
dead leaves during the time the moon is visible. Itis a vegetarian, 
and in search of food often climbs shrubs and small trees. A great 
article of food with the natives.”—C. H. B. G. 

3. 1569 GQmmature). 

‘“‘ Native name, ‘ Masengwi.’ 

‘‘ Not very common, and apparently somewhat locally distri- 
buted. Habits similar to its congeners in other parts of 8. Africa, 
forming the usual burrows in the more open forests, never in 
the thickets where Cricetomys gambianus is usually observed. 
They were extremely wary and difficult to trap and bad luck was 
experienced with them, inasmuch as in several instances only the 
toes were left in the trap and in one case the black end of the 
tail. ‘ Majengwi’ was the name given to this species by the natives 
at the Klein Letaba, hence this isan added proof of the existence 
of the Springhaas in that loeality.”—C. H. B. G. 


3. 1518, 1542, 1563, 1565. ©. 1568, 1667, 1669. 

A Hare of the capensis group with the nuchal patch grey, and 
the chin, chest, and belly snowy white. 

Fur long, soft, and very fine, length on middle of back about 
25 mm. General colour of upper surface “ broccoli-brown ” 
tinged with ‘“drab-grey,” becoming lighter and gradually suffused 
with butty on the flanks. Individual hairs divided into five rings 
of the following colours and approximate dimensions :—proximal 
ring “grey no. 9,” 7 mm. in length; second ring very pale ecru- 
dvab, 5mm.; third ring jetty black, 4 mm.; fourth ring light 
pinkish buff, 6 mm.; distal ring black, 3mm. Individual hairs 
of belly about 30 mm. in length, soft, silky, and entirely snow- 
white. Nostrils and lips edged with white, muzzle and vibrissate 
area between wood-brown and isabella colour; a broad line ex- 
tending from upper edge of orbit to within half an inch of muzzle, 
and thence spreading downwards over lower region of cheek sandy 
ervey; infraorbital area between wood-brown and isabella colour. 
Eyes ringed with white; forehead coloured like back. Ears dis- 
tinctly smaller than in the other subspecies, with the proectote* 
coloured like forehead and back, fringed with yellowish-buff hairs 
intermixed with white; metentote sparsely covered with minute 
white hairs, outer fringe snowy white, absent terminally ; 
metectote white, the terminal margin black. Nape-patch between 
“ smoke-grey”” and “drab-grey.” Interramia white. Throat- 
patch “pinkish buff.” Chest and belly snowy white. Outer side 
of fore and hind limbs deep buff, becoming lighter and interspersed 
with white on fore and hind feet. Tail short, compactly haired, 
not loose and straggling as is generally the case in specimens of 
ochropus, black above, white below. 

Skull as in true capensis. 

Dimensions of the type (measured in the flesh) :— 

Head and body 472 mm.; tail 95; hind foot 113; ear 92. 

Skull—greatest length 85°5 mm. ; basilar length 70; zygomatic 
breadth 40-1; nasals, oblique length 38:2; brain-case breadth 
28°0; diastema 24-4; palatal length 36-0; palatal foramina 22-7 ; 
upper molar series 14:7; antero-posterior diameter of bulla 11:3. 

Type. Male. B.M. ne. Original number 1565. 
Collected 6 July, 1906. 

This subspecies may be distinguished from Z. ochropus by its 
white chin and chest and the buffy colour of its flanks, in contrast 
with the yellow of the latter on the sides and nape. It can be 
easily separated from Lepus capensis centralis and L. c. granti by 
their having a vinaceous buffy or pinkish buffy chest and flanks. 
It bears more superficial resemblance to ZL. c. capensis however, 
but is distinguishable by having a white chest and chin and a 
white border surrounding the tip of the ear, instead of, as is usual, 

* Thomas, P. Z. S. 1905, vol. ii. p. 359 (1906). 


the terminal portion edged with black. A comparison with a 
considerable series of all the forms mentioned above shows that 
examples of the present subspecies have a considerably shorter 
tail and ears than is usual in any of them. Further material will 
probably show that intergradation takes place between this form 
and Z. ochropus, in which case the latter will take its place in 
the group as a subspecies. 

‘“¢ Native name, ‘ Nfundla.’ 

‘‘Gommon everywhere, especially in the valley of the Inyasuni. 
Generally lying up during the daytime in the clumps of small 
bush or grass, and feeding throughout the night, when they can 
be taken with a noose on the numerous footpaths. Numbers are 
caught by the natives in this way, the animal being with them a 
staple article of food.”—C. H. B. G. 

Q. 1870. 

Tn its strong ferruginous colouring this specimen quite agrees 
with examples from Zomba, B. C. Africa, to which Dr. Forsyth 
Major has given (P. Z.8. 1897, p. 367) the subspecific name nyase. 

“Native name, ‘ Ngulubi.’ 

‘Plentiful in the thickets and dense forests, but extremely 
difficult to secure. Always observed in pairs, and more or less 
nocturnal in habits. _When pursued, they savagely attack the 
dogs, repeatedly charging until killed by the hunters.”—C. H. B.G. 


3. 1613, @. 1485, 1675. 

‘“¢ Native name, ‘ Mungulwi’ or ‘ Munguli.’ 

‘‘Not common, and found in the dense forest and thickets, 
which the ‘ Nhlengane’ loves; even there it is locally distributed, 
seeming to confine itself to certain patches, from which it never 
wanders far. Only one or two others, besides the specimen sent, 
were observed, they were not however secured.”—C. H. B. G, 


@. 1503, 1582, 1640. 

‘“‘ Native name, ‘ Munti.’ 

‘“¢ Not too common, and found both in the river-valleys and the 
forests, visiting the Kaffir lands at night. Numbers of this and 
other buck are taken by the natives for food.”—C, H. B. G. 


Ss WL, USGA, Sy Ibeyet) 

These specimens quite agree with those from Klein Letaba, 
N.E. Transvaal. 

‘““ Native name, ‘ Isipenu.’ 

“ Fairly plentiful but locally distributed, confining itself to the 
more open forest and plain, along the river-valleys, away from 
habitations. Generally observed in pairs.”—C. H. B. G,. 



@. 162 Wore) 9% 15255 1576; 16225 

The examination of these specimens makes it clear that the 
subspecies zulwensis Thos. of the Nyasan livingstonianus is quite 
constant in the essential characters in which it differs from the 
latter, and we think ourselves justified in regarding zulwensis as a 
distinct species. 

‘“‘ Native name, ‘ Nhlengane.’ 

“Very common at Coguno and the whole country inland, but 
unknown in the immediate neighbourhood of Inhambane. It 
inhabits the dense thickets and undergrowth, where it has regular 
tracks, and is generally observed in twos, and occasionally threes. 
It can be easily obtained in the early mornings and late afternoons, 
when it is found out feeding in the more open patches of bush 
and along the Kaffir footpaths that intersect the thickets in every 
direction, or by joining a hunt which the natives organise for the 
special purpose of taking this species for food. It is an extremely 
difficult buck to see when standing in the thickets, and can often 
be heard giving a goat-like snort of alarm, although quite invisible 
at only a few yards distance. The does greatly outnumber the 
bucks, and it is impossible to make out the sexes in their habitat ; 
a great proportion of the former are killed, and very old examples 
of the latter are not often obtained.”—C. H. B. G. 



“‘ Native name, ‘ Mhlangu,’ 

“Observed only in the river-valleys, in many of which they 
are very plentiful, often being seen six and eight together. In 
the valley of the Inyasuni, where they are little disturbed, they 
feed throughout the day, and are remarkably tame. Individual 
males vary much in the thickness of the neck.”—C. H. B. G. 

April 9; 1907: 
Dr. Heyry Woopwarp, F.R.S., Vice-President, in the Chair. « 

Mr. RK. I. Pocock, the Superintendent of the Gardens, exhibited 
a photograph and the skull of a specimen of the Manul or Pallas’s 
Cat (Felis manul) that had recently died in the Society’s Menagerie, 
and made some remarks on the species. 

The specimen (text-fig. 87) was received in exchange from the 
Zoological Gardens in Calcutta, in April 1906, and died from 
broncho-pneumonia in April 1907. It was alleged to have come 
from Tibet. Untortunately no exact locality was recorded; but 
since the coloration of the skin and the structure of the skull 
agree with those of examples from Tibet that have been described, 

300 MR. R. I. POCOCK ON PALLAS’S CAT. [ Apr. 9, 

there is no reason to doubt that the animal was captured in that 
country *. 

When alive, this Cat differed markedly, both in behaviour 
and appearance, from most captive examples of the genus /elis. 
The latter, if they happen to be tame, usually evince grati- 
fication of notice by rubbing, with tail erect, against the bars 
of the cage. When wild, they are either contemptuously in- 
different to friendly overtures or receive them, crouched in a 
corner, snarling. The Manul, on the contrary, although not tame 
enough to be handled or touched, nor sufficiently friendly to rub 
himself against the bars, showed no fear of spectators and no 
wish to avoid them. He would boldly and aggressively but 
silently advance to the front of the cage and, standing on his hind 
legs, grip the bars with his fore paws, ready to scratch at a 
confiding hand unwarily placed within reach. 

Text-fig. 87. 

Felis manul. 

(From a photograph of a specimen living in the Society's Gardens.) 

Like all the small Cats, he was usually very silent. He was 
never heard to utter the growling snarl and guttural expiratory 
hiss with open mouth so familiarly associated with irritation 

* Except that the orbits are incomplete behind, the skull closely resembles that of 
a speciinen from Ladak in the British Museum. 


1907. | MR. R. I. POCOCK ON PALLAS’S CAT. 301 

of temper im Cats; and the “spit” was a short, sharp sound 
like “ts,” “ts,” “ts,” projected through nearly closed lips. 
According to the keeper, Dixon, the “mew” or “ caterwaul ” 
was a sound somewhat recalling a combination of the bark of a 
small dog and the “hoot” of an owl. This was heard on one 
occasion in reply to the typical “ caterwaul” of a female Uganda 
Cat (/. ocreata), when “on heat.” 

The tail was almost invariably carried with its posterior half 
upcurled, so that the broad black confluent stripes on the under 
side of its distal end were in full view from behind. The 
conspicuousness of this jet-black area was enhanced in a marked 
degree by his peculiar habit of jerking the end of the tail smartly 
up and down. The lowness and width of the summit of the head 
and the lateral “set” of the ears imparted to this Cat an aspect 
totally different from that of all other species (text-fig. 87). In the 
latter the inner edge of the ear normally rises obliquely outwards 
from the top of the head to form, with the outer border, a con- 
tinuous curve or anacute angle. Butin /’. manul the inner border 
lies normally in the same plane as the top of the head, and meets 
the vertical outer border at a right angle. This border rises from 
the head at a point on a level with the outer canthus of the eye. 
Since depression of the ears in Cats is an infallible sign of anger 
or of predatory excitement, the simulation of this act caused by 
the low and lateral setting of these organs in the Manul imparts 
to his face a permanent look of ferocity and unrest, quite unlike 
the placid aspect of other Cats with their ears normally erect. 

So far as the Manul is concerned, one practical result of the 
lowness of the forehead and the lateral setting of the small ears 
is the power to peer over the edge of an object, like a rock or a 
fallen tree-trunk, without depressing and closing the ears, and 
without showing so much of the cranium as most of the “ high- 
headed ” Cats do when so occupied. Other Cats, when watching 
prey from behind some such point of vantage, always lower the 
ears so as to make them invisible and, at the same time, partially 
close them in such a way that quickness of hearing must be 
interfered with to a greater or less extent. In this, perhaps, 
may be found the explanation of the peculiar structural features 
in the head which give the Manul its remarkable physiognomy. 
Be this as it may, the above-proffered explanation was forcibly 
suggested by the observation of the living Manul peering over 
the edge of his sleeping-box and showing a relatively small 
enon of head above the eyes, the ears at the time being 
scarcely perceptibly depressed and not in any sense closed. 

It has been stated by Gray *, on the authority of Hodgson, 
that the pupils of the eyes are linear and erect. This was not 
the case in the Society’s specimen. Under the influence of 
sunlight the pupil contracted to a small circular or subcircular disk. 
The ivis was yellowish. 

* P. Z.S. 1867, p. 275. This statement probably misled Elliot into having the 
eyes of F. manul in his monograph drawn like those of a domestic Cat. 

Proc, Zoou. Soc.—1907, No. X XI. 21 

302 MR. R. I, POCOCK ON PALLAS’S CAT. [ Apr. 9, 

There was no marked seasonal change in colour, the coat 
merely becoming thicker in winter than in summer. 

Satunin has recently made Felis manul the type of a new 
genus, Zrichelurus (Ann. Mus. St. Pétersb. ix. pp. 495-506, 
1905), being the first to point out in detail the structural 
peculiarities of the species. Unfortunately he overlooked the 
fact that Severtzow had already proposed the name Otocolobus for 
the same species (Rev. Mag. Zool. x. p. 386, 1858). 

To Satunin also belongs the credit of showing that three 
distinct. forms of Manul are recognisable, each typical of a par- 
ticular geographical area. To two of these he gives subspecific 
and to one specific rank. 

Substituting Otocolobus for Trichelurus, his classification is as 
follows :— 

la. Otocolobus manul Pall. (typical form), from Trans- 
caspia, Turkestan, and Siberia to the west of Lake 
b. Otocolobus manul mongolicus Sat., from Mongolia and 
Siberia to the east of Lake Baikal. 
2. Otocolobus nigripectus Hodgs., Tibet. 

I have not seen skins of the typical or of the Mongolian forms, 
but, judging from Satunin’s description and the published figures, 
I should say that the differences between them and the Tibetan 
form are only of subspecific value. The name for the latter 
therefore will be Otocolobus manul nigripectus. 

The Society’s specimen of this subspecies presented the following 
characters :— 

Prevailing colour of face grey: some buff below and at the 
sides of the nose: the eyes surrounded by a greyish-white area 
which is bordered bya black streak above and below and partially 
on the inner side, giving a characteristic spectacled look to the 
face. The buff area round the nose set off by a black patch, 
whence arise some black moustachial bristles; the rest of these 
bristles mostly white and arising from blackish lines on the 
whitish area of the upper lips. Some black spots running into 
abbreviated lines on the cheek below the eyes. The two genal 
stripes broad and jet-black, descending obliquely downwards and 
backwards, the inferior arising from a spot on a level with the 
middle of the eye, the superior from a point near the outer canthus 
of the eye; area above the latter stripe grey, below the former 
pale grey turning to white posteriorly ; the area between them 
pale greyish white. Posteriorly the two stripes are confluent, 
and merge below the ear with the sooty-brown hue of the throat 
and chest. The dark hue of these areas relieved by the long 
white tips to the hairs. Summit of head black speckled with 
white; its fore part or the area in front of the ears marked with 
some small asymmetrically disposed jet-black spots interspersed 
with a few greyish-white spots. Back of ears greyish, passing 

1907. | MR. R. I. POCOCK ON PALLAS’S CAT, 303 

into black towards the head and with a darker rim; hairs on the 
inner side and in front of the ears whitish grey. Prevailing hue 
of the dorsal area of the neck and body silvery- or iron-grey, the 
hairs sooty black basally, then white witha black tip*. Laterally 
the dark-tinted basal portion of the hairs is paler in hue and less 
in extent, the greater portion of the hair below the white sub- 
apical band being whitish and strongly or faintly tinged with buff, 
the buff tint showing up strongly when the hairs are parted. On 
the lower side of the body the hairs have no black tint and the 
buff tint disappears, leaving the belly and chest white but for a 
clouding of blackish blotches on the chest behind and between the 
fore legs. On the lumbar and sacral regions of the back there are 
traces, mostly very faint, of narrow transverse black stripes. 
The largest and most distinct of these lies about midway between 
the shoulder and the root of the tail; on the right side it 
measures about 57 mm., on the left about 50, the two being 
separated by a median area of about 20mm. ‘This is the only 
stripe that is evident on the right side. On the left side there is 
one short stripe in front of it, and two abbreviated stripes behind 
it, one low down on the side, the other higher up and just 
traceable to the middle line. Behind this there are, on the 
middle line of the back, three faint transverse blackish blotches, 
probably representing the dorsal end of stripes. The tail is 
greyer and paler than the back ; in its distal half there are three 
well-marked black stripes, narrow dorsally and laterally, but 
expanding and forming a triangular patch inferiorly where they 
meet. The last stripe fuses laterally and inferiorly with the 
terminal black tip; and since the expanded areas of the other 
stripes are only separated by the narrowest intervals, it follows 
that the distal half of the tail is practically black below. The 
proximal half of the tail is marked above with three narrow 
indistinct stripes, which, however, widen and become much more 
strongly defined below. Fore leg whitish grey turning to creamy 
buff on the paws; the brachial stripe distinct: other stripes 
represented by blackish patches. Hind legs with thighs grey 
like the body, and indistinctly spotted ; front of the leg greyish, 
turning to pale buff on the paws; back of the leg up to the 
hock rusty cream, a black patch on each side of the paw above 
the median pad. 

Measurement of dressed and flat skin :—Head and body 20 inches 
(=500 mm.), tail 9 inches (= 225 mm.). 

It is noticeable that the hair of the ventral surface, both on 
the throat, chest, belly, and thighs, is considerably longer than on 
the dorsal and lateral surfaces of the body. On the spinal region 
it measures about 23 mm., and on the belly about 43 mm. The 
coating of long hairs below is perhaps an adaptation to life in 

* Satunin (op. cit. p. 497) says that the hairs on the back and sides have white tips. 
Did he overlook the slender black tip ? 

304 MR. R. I. POCOCK ON PALLAS’S CAT. | Apr. 9, 

Text-fig 88. 


Skull of Felis manul, viewed from above. Nat. size. 

cold latitudes, where lying or sleeping in the snow is possibly not 
an uncommon occurrence, In that case the hairs would act as a 
protection against chill to the delicate internal organs, especially 
the intestinal portion of the alimentary canal *. 

The skull of 7. manul has been described briefly by Milne- 
Edwards and Blanfordt, and more in detail by Satunin, who gives 
measurements of three examples. The skull of the specimen that 
lived in the Gardens (text-figs. 88, 89) does not apparently differ 
greatly from these. The chief peculiarities of the skull may be 
shown by comparing it with skulls of Felis sylvestris and Felis 
ocreata, since £, manwl is, im my opinion, an aberrant form of 
the group exemplified by these two species. 

* The ventral development of the hair in the Yak (Bos grunniens), also a denizen 
of cold countries, is a parallel case. The size of the tutt of hair at the end of the 
tail inthis animal perhaps acts as a protection against frost-bite of a part of an organ 
where the circulation is weakest. 

+ Rech. Mamm. p. 226. ¢ Mamm. Brit. India, p. 83. 


Text-fig. 89. 

Skull of Felis manul, side view. 

Width across zygomata ......... 

behind postorbital processes ... 
Ot: DLAI CARL ae ene up arabian 
lessee DELYVEEMPOLDITS ye eisen er: 

», of mesopterygoid fossa 
Length of palate .......... 


PT ROLEZ UE OMe Crate en eeyenee ree 

Height from coronoid to angular process 
Length of upper carnassial 

PP OL LO Wer CALNASsIAl ones enes 
Median length of nasals ......... 

distal edge 

just above constriction 

Shope! Merayeisler Ose jSebUUl | iaedsanopngoon panos nondon 
Jebyse yl Memmye*sioi Coe SVC a jenn en oogbed Indnad asuro anal 

across postorbital processes ............ 

| ., ot muzzle at base of canines ............ 
» of muzzle at infraorbital foramina 

| ,, between auditory orifices Ae 
») ©ACKOSS Uppel Ccarnassials’.................. 

from palate to occipital foramen .... 

» Of mandible from condyle............... 

Width of nasals in line with transverse 

across constriction ......... 

FE. manul EF. sylvestris 


67 | 
69 | 
A7 | 
19 | 
Py | 
23 | 





U2? Mh 
EAH ay 



| FP. ocreata | 

i belay 
50 | 

306 MR. R. I, POCOCK ON PALLAS’S CAT. [ Apr. 9, 

Some of the principal differences not shown in the above-given 
measurements may be described as follows :— 

é. Upper carnassial without inner lobe; no maxillary excres- 
cence bounding the infraorbital foramen above and out- 
side; malar sending up a long narrow process in front of 
and considerably above the lacrymal foramen; infero- 
anterior edge of orbit circularly rounded ; upper edge of 
orbit elevated, higher than median portion of frontal bone ; 
facial portion of skull abruptly inclined; occipital crest 
small; interparietal and parietal crests absent ; temporal 
crests marked by two shallow grooves nearly evenly 
converging from the postorbital processes to the inter- 
parietal; the smooth median and rougher lateral (tem- 
poral) area of the parietals slightly elevated and separated 
by a shallow depression ; fronto-parietal suture strongly 
angled behind the postorbital processes and procurved 
(concave forwards) dorsally ; basisphenoid longitudinally 
BMCHOG RA as feratetanr ou caciben be cia eRe eran eer aces manul. 

b. Upper carnassial with strong inner lobe; a distinct excres- 
cence on the maxilla bounding the infraorbital foramen 
above and externally ; upper process of malar not extending 
past the lacrymal foramen ; infero-anterior edge of orbit 
more ovally rounded ; upper edge of orbit depressed, lower 
than median portion of frontal bone; facial portion of 
skull evenly curved downwards from the frontal to the 
middle of the nasals; occipital and interparietal crests 
well-developed ; temporal crests, when not mesially con- 
fluent, represented by a low ridge defining a median lyrate 
area forming a practically continuous curve with the 
temporal portion of the parietal bone; fronto-parietal 
suture forming a nearly straight transverse line; _basi- 
sphenoid nearly flat, only lightly arched longitudinally. 

sylvestris and ocreata. 

The skulls of two Tibetan specimens that I have seen do not 
agree with Milne-Edwards’s statement that the nasals in Mongolian 
specimens are strongly compressed in their posterior and corre- 
spondingly dilated in their anterior portion. Satunin also states 
that the nasals in the Tibet Manul (/. nigripectus) differ from 
those of the typical form in being constricted in the middle and 
distally expanded. In the skull of the animal in the Society’s 
Collection, the constriction is very slight, amounting to only | mm. 
in transverse width across the two bones. From this constriction 
the bones expand very gradually forwards and backwards. ‘They 
may be described as being of the broad type, such as is shown in 
F’, sylvestris. 

The infraorbital foramen is small and vertically oval, its 
greatest length being less than the distance between its upper 
extremity and the superjacent edge of the orbit. 

iw Ha 



“GOlIaS ae “1lOnhea sheave l 

“duit uesan'p UAT TSP elseeg YW 



‘duit wsex) “Pp 


“UAT esp eTTeeS 

let Ww 


The following papers were read :— 

1. On a small Collection of Fishes made in the Eastern 
Watershed of the Transvaal by Capt. G. H. Bruce. 
By G. A. Bounencsr, F.R.S., F.Z.8. 

[ Received March 12, 1907. ] 
(Plates XVIII. & XIX., and Text-figures 90, 91.) 

When recently serving in the Transvaal, Capt. G. E. Bruce, 
5th Mounted Infantry, was so kind as to respond to an appli- 
cation I made to him in 1905, to preserve some of the fishes. 
in which he was interested, and which I felt sure would prove 
of considerable scientific value, so little being known of the 
piscine fauna of the Transvaal. I have now the pleasure of 
drawing up a list of the species represented in a small collection 
made by Capt. Bruce and presented by him to the British 

The rugged nature and muddy water of the Transvaal rivers, 
together with the steep banks overgrown by reeds and rushes, 
make it very difficult to collect, except by fishing with fly or bait, 
and the laws against netting and dynamite ase very strict. 
Further, the size of the collecting-jars at Capt. Bruce’s disposal 
precluded the preserving of large specimens. Notwithstanding 
these restrictions, the collection contains examples of several 
species not previously recorded from the Transvaal, and of five 
which are here described as new. 


1. Hyprocyon LinzeAtus Blkr. 

The ‘“ Tiger Fish” occurs in the Inkomati and Krokodil Rivers, 
and appears to be plentiful in all the rivers of the East Coast. 
from the Zambesi to Swaziland. 


New to the Transvaal, where it was found in the Inkomati 
River at Komati Poort. First described from the Zambesi, it. 
has since been found in Lake Nyasa, in German East Africa, in 
the Congo, and in Angola. 

Inkomati River at Komati Poort. Not previously known 
from south of the Zambesi. 

Three specimens from Groot Olifant River agree well with 
Steindachner’s description and figure of a specimen from the 


Limpopo River, except for the presence of two pairs of barbels. 
I cannot help thinking that the absence of anterior barbels in 
the type specimen is due to an anomaly, or that their presence 
has been overlooked. 

5. Barus EuTzNIA Bler. 

Several specimens from Klein Olifant River. This species, 
originally described from Angola (B. kessleri Gthr. nec Stdr.), has 
recently been rediscovered in N.W. Rhodesia by Mr. Neave. 

6. BARBUS POLYLEPIS, sp. n. (Text-fig. 90.) 

Depth of body 41 times in total length, length of head 33 times. 
Snout rounded-subacuminate, 3 times in length of head, feebly 
projecting beyond mouth ; diameter of eye 41 times in length of 
head, interorbital width 33 times; mouth small, inferior, its 
width 4 times in length of head; lips well developed, lower 
continuous across chin; two barbels on each side, subequal in 
length, 3 diameter of eye. Dorsal III 8, last simple ray strong, 


bony, not serrated, its rigid part # length of head; free edge of 

Text-fig. 90. 

——— S CO 


Barbus polylepis. 

the fin strongly emarginate; its distance from posterior border 
of eye equals its distance from caudal. Anal IIT 5, longest ray 
3 length of head, not reaching root of caudal. Pectoral 3 length 
of head, not reaching ventral; latter below anterior rays of 
dorsal. Caudal deeply forked, with pointed lobes. Caudal 

peduncle twice as long as deep. Scales 43 oh 5 between lateral 
Ime and ventral, 18 round caudal peduncle. Olive-grey above, 
white beneath. 

Total length 120 millim. 

A single specimen from Klein Olifant River. 

In the Groot Olifant River, Capt. Bruce obtained a large Barbus, 
weighing 32 lbs., and too large for him to preserve, which, from 
the notes he has taken, probably represented the adult of the fish 
here described. Capt. Bruce counted 42 scales in the lateral line, 
84 in a transverse series above the lateral line, and 6 between 
the latter and the ventral fin. This large specimen further 
differed in the much longer rays of the anal fin. 



Groot Olifant River. 
This fish may be the same as B. natalensis Castelnau, in- 
sufficently described. 

8. BARBUS BRUCH, sp. n. (Plate XVIII. fig. 1.) 

Depth of body 33 times in total length, length of head 4 times. 
Snout rounded-subacuminate, 2? times in length of head, strongly 
projecting beyond mouth ; diameter of eye 5 times in length of 
head, interorbital width nearly 3 times; mouth small, inferior, 
its width 43 times in length of head; lips strongly developed, 
lower continuous and forming a rounded mental lobe; two 
barbels on each side, anterior not quite 3 diameter of eye, 
posterior 3. Dorsal IV 9, last simple ray strong, bony, not 
serrated, its rigid part 3 length of head; free edge of the fin 
strongly emarginate; its distance from centre of eye equals its 
distance from caudal. Anal III 5, longest ray 2 length of head, 
not reaching root of caudal. Pectoral ? length of head, not 
reaching ventral; latter below middle of base of dorsal. Caudal 
deeply forked, with poimted lobes. Caudal peduncle once and 

% as long as broad. Scales 29 z, 2 between lateral line and 
ventral, 12 round caudal peduncle. “ Muddy grey above, white 
beneath; dorsal yellowish grey; caudal and paired fins bright 

Total length 150 millim. 

A single specimen from the Groot Olifant River. 

9. Barsus sEcTOR, sp.n. (Plate XVIII. fig. 2.) 

Depth of body 33 times in total length, length of head 44 times. 
Snout rounded, 3 times in length of head, strongly projecting 
beyond mouth ; diameter of eye 4 times in length of head, inter- 
orbital width 2? times; mouth inferior, feebly curved, its width 
3 times in length of head, lower jaw with a sharp edge; lower 
lip restricted to the sides; two barbels on each side, anterior 
about 3, posterior about 4 diameter of eye.. Dorsal IV 9, last 
simple ray strong, bony, not serrated, its rigid part 2 length of 
head ; free edge of the fin feebly emarginate ; its distance from 
centre of eye equals its distance from caudal. Anal III 5, 
longest ray # length of head, not reaching root of caudal. 
Pectoral a little shorter than head, not reaching ventral; latter 
below middle of base of dorsal. Caudal deeply forked, with 

pointed lobes. Caudal peduncle once and # as long as deep. 
Scales 29 Z, 23 between lateral line and ventral, 12 round caudal 
peduncle. “ Muddy grey above, white beneath ; dorsal yellowish 
grey; caudal and paired fins bright pink.” 

Total length 140 millim. 

A single specimen from the Groot Olifant River. 

But for the shape of the head and the structure of the mouth, 
owing to which it should be referred to the genus Capoéta of 


Giinther, this fish could not be distinguished from the preceding. 
Similar cases of close resemblance between species thus referable 
to Barbus and Capoéta have been pointed out by me when 
describing collections from Morocco and Kast Africa. 

10. BARBUS ELEPHANTIS, sp.n. (Text-fig. 91.) 

Depth of body equal to length of head, 4 times in total length. 
Snout rounded, 3 times in length of head, projecting beyond 
mouth; diameter of eye 4 times in length of head, interorbital 
width twice and 3; mouth inferior, feebly curved, its width 
33 times in length of head, lower jaw with a sharp edge; lower 
lip restricted to the sides; two barbels on each side, anterior 
= diameter of eye, posterior as long as eye. Dorsal IV 8, last 
simple ray strong, bony, not serrated, its rigid part 3 length of 

Text-fig. 91. 

Barbus elephantis. 

head; free edge of the fin strongly emarginate; its distance 
from posterior border of eye equals its distance from caudal. 
Anal ITT 5, longest ray 2 length of head, not reaching root of 
caudal. Pectoral ? length of head, not reaching ventral ; latter 
below middle of base of dorsal. Caudal deeply forked, with 


pointed lobes. Caudal peduncle once and % as long as deep. 
Scales 37 2 3 between lateral line and ventral, 14 round caudal 
peduncle. ‘‘ Muddy grey above, white beneath; dorsal yellowish 
grey ; caudal and paired fins bright pink.” 

Total length 155 millim. 

A single specimen from Groot Olifant River. 

Very similar to the preceding species, but distinguished by longer 
barbels and smaller scales. 

11, Barus trimacutatus Peters. 

Groot and Klein Olifant Rivers and Inkomati River. 

12. Barsus INERMIs Peters. 

Klein Olifant River. . 

Previously known from the Zambesi only. 

13. VARICORHINUS BRUCI, sp. n. (Plate XIX.) 

Body strongly compressed, its depth 33 times in total length ; 


length of head 42 times in total length. Snout rounded, broader 
than long, 4 length of head; eye lateral, 43 times in length of 
head, twice in interorbital width; no conical tubercles on the 
head ; mouth feebly curved, its width 3 times in length of head ; 
two barbels on each side, anterior 4 diameter of eye, posterior as 
long as eye. Dorsal TV 9, last simple ray strong, bony, not 
serrated, its rigid part 3 length of head; border of fin concave ; 
longest ray a little shorter than head. Anal IIT 5, not reaching 
root of caudal. Pectoral a little shorter than head, not reaching 
ventral, which is inserted below middle of dorsal. Caudal 
deeply forked, with pointed lobes. Caudal peduncle twice as long 
as deep. Scales 31 a 24 between lateral line and ventral, 
12 round caudal peduncle. “ Back muddy brown, belly white, 
fins grey.” 

Total length 170 millim. 

A single specimen from Klein Olifant River. ‘“ Lives in deep 
still pools or under big rocks.” 

Closely allied to the recently described V. ansorgii Blgr., from 
the Kwango River (Congo System) in Angola, which has a deeper 
body, a shorter caudal peduncle, and the eyes turned more 
upwards. This is the fifth known African species of Varicorhinus. 

Groot Olifant River. 

Groot Olifant River. The specimen is referable to 4. labiata 

Groot Olifant River. 

Inkomati River, near Komati Poort. 

Groot and Klein Olifant Rivers. 


Pratt XVIII. 

Fig. 1. Barbus brucii, p. 309. 
2. ar sector, p. 309. 
a. Mouth seen from below. 

PuatEe XIX. 

Varicorhinus bructi, p. 310, with upper view of head and mouth 
seen from below. 

312 MR. T. A. COWARD ON THE [| Apr. 9 

On the Winter Habits of the Greater Horseshoe, Rhino- 
lophus jferrum-equinum (Schreber), and other Caye- 

haunting Bats. By T. A. Cowarp, F.Z.8. 
[Received February 7, 1907. ] 

Jn my paper “On some Habits of the Lesser Horseshoe Bat, 
thinolophus hipposiderus (Bechstein)” (1) I referred to several 
points of interest in connection with that species during its 
winter retreat, and suggested that its congener, the Greater 
Horseshoe Bat, Rhinolophus ferrum-equinum (Schreber), would 
be found to have similar habits. Since writing that paper I 
have had the opportunity of observing the Gr eater Horseshoe in 
winter in caves in the Mendips, and am fortunate in being able to 
compare my notes with those made by Mr. Bruce F. Cummings, 
of Barnstaple, Devon, who at the same time was making 
observations on the species in the neighbourhood of that town. 
The conclusions that I arrived at require some modification, but 
only one of them was not fully confirmed: I said that the bats 
“usually” occupy different retreats in summer and winter; it would 
be more correct to say that at times and in certain localities different 
retreats are used at different seasons, for in the Cheddar caves the 
bats occupy apparently similar positions in summer and winter. 
I have, indeed, little knowledge of the habits of the bats in this 
locality during the summer months, but I found the remains of 
insects which must have been captured in summer lying beneath 
places which the bats occupy in winter. 

My observations were made at Cheddar, Somerset, and in the 
neighbourhood, between December 27th, 1906, and January 7th, 
1907, and, later, on a few examples in captivity. Mr. Cummings 
paid a number of visits to some disused manganese mines in the 
Pickwell Down Sandstone near Barnstaple between December 24th, 
1906, and January 23rd, 1907, and also kept one bat alive for some 
days. At the end of December and beginning of January I 
found Greater Horseshoes both in the deepest recesses of the 
caves—some of which are of great length—and close to the 
entrances, even within reach of daylight. 

I regret that I have no accurate record of the temperature in 
the open at Cheddar during my visit, but on the first few nights 
there were sharp frosts; later the weather was mild and at 
entrances of the caves the temperature varied from 40° to 43° F. 
Within the caves—at some distance from the entrances—the 
temperature was fairly constant; in different spots [ found it 
between 50° and 52° F., and this I understand is also the summer 
temperature. Mr, Cummings also found that the temperature in 
the Barnstaple mines was 52° 10 

The dimensions of the caves which I visited vary considerably. 
The Long Hole, in which JI found the Greater Horseshoe 
abundant, is a long natural tunnel in the limestone of perhaps 150 
or 200 yards in length, and with an average width of seven or 


eight yards and a height of from five to ten feet; in places the 
roof is much higher and culminates in fissures and hollows which 
afford suitable hiding-places for large numbers of bats. The floor 
is uneven and partially blocked by ancient roof-falls. At the end, 
beyond a considerable mass of fallen rock, the cavern opens into 
a small chamber, from which a chimney, blocked by a huge “‘ chock 
stone,” runs upward to a chamber of considerable size. Out of 
this upper chamber a low passage, in places less than eighteen 
inches high, extends for some yards; the end of this passsage 1s 
blocked, and when I examined it no flicker was noticeable in the 
flame of my candle, but [ was informed that at one time it was 
possible to feel a current of air near its termination, as if it led to 
some outlet. From the lower chamber a lime-encrusted shaft, 
running upwards, may lead to further chambers or to the open. 
During a portion of my visit I was accompanied by Mr. Charles 

In the Long Hole I found Greater Horseshoes occupying a 
fissure high up in the roof near the entrance. I cannot say how 
far within this fissure the bats rested in the daytime, for they 
were not visible; it was only after dusk that I saw them emerge 
from some hole in the fissure. I found other Greater Horseshoes 
scattered singly in the main tunnel, hanging from the roof or in 
the water-worn holes with which it is abundantly pitted. In 
the terminal chamber the bats were in colonies of varying size. 

In one of the caves of the village, which is open for visitors and 
illuminated by gas, there is, near the entrance, a wide shaft open 
to the air. Some ten feet from the floor of the cave, and perhaps 
fifteen feet from the top of the shaft, a small side-chamber opens 
out of the wall; the floor of this chamber was thickly covered with 
old dry dung, mingled with fragments of many beetles, chiefly of 
the genus Geotrupes. Dr. J. Harold Bailey, who kindly examined 
all my coleopterous fragments, identified portions of Geotrupes 
spiniger Marsh, and of another Geotrupes. The Greater Horse- 
shoe, as I shall show later, captures some, and in summer probably 
all, of its prey outside the caves and conveys it to the interior of 
the caves before consuming it. The head, elytra, and other 
portions of the beetles are dropped whilst the bat is crushing the 
insect in its jaws. his is undoubtedly a common habit, but bats 
which go far from the caves in search of food may hang to 
convenient footholds at some distance from their diurnal sleeping- 
places. In captivity the bats rest after a meal and frequently 
defecate, a habit which probably accounts for the piles of dung 
beneath favourite feeding-places in the caves. Both Mr. Cummings 
and I noticed that our captive bats showed a marked preference 
for certain resting-places, settling again and again within a few 
inches of the same spot. This chamber in the shaft was perhaps 
the only place I came across which appeared to be solely a summer 
haunt; it may or may not be occupied as a sleeping-place in 
summer, but it was undoubtedly a chamber to which numbers 
retired to feed. 

314 MR. T. A. COWARD ON THE [Apr. 9, 

Great Oons Cave is longer than the Long Hole; it consists of 
a rough, slightly descending tunnel and a steep passage running 
downwards to a considerable depth from the end of the main tunnel. 
At the bottom of this terminal passage, at a great distance below the 
entrance of the cave, I found one Greater Horseshoe hanging ; 
a few others and Lesser Horseshoes, Rhinolophus hipposiderus 
(Bechstein), were in other parts of the cave, and a Whiskered 
Bat, Myotis mystacinus (Leisler), was close to the entrance. The 
Kcho Cave is a great cleft in the cliff, entered by a small hole and 
short passage; the cleft then extends upwards and downwards, the 
height being over a hundred feet and the width little more than 
eight feet. Thirty feet or more above the floor of the cave 
colonies of Greater Horseshoes were hanging from the wall on 
one side, and a few Lesser Horseshoes were suspended, singly, 
from the opposite wall. Goatchurch Cavern, on the northern 
slope of the Mendips, has its entrance high on the hill-side in the 
treeless valley of Burrington Combe. This cavern consists of long 
and intricate passages, running far into the hill and descending 
steeply; here again at a great depth L found both species of 
Horseshoe scattered about the passages. In other smaller and 
shallower caves, some with narrow entrances and others with wide 
mouths, I found Greater and Lesser Horseshoes; one of the 
former hung in full daylight ten yards from the entrance, and a 
Lesser Horseshoe from the upper part of a bottle-shaped hollow 
in the roof only four yards from the entrance. 

In no ease in which I was able to make repeated observations 
were the bats actually hibernating. All were susceptible to the 
disturbance caused by my entrance, drawing themselves up by 
bending their legs and often swaying slightly, although not 
touched ; bats changed their positions in the intervals between my 
visits, while others, as I shall show later, flew in the caves 
without any artificial stimulus and indeed went out into the open 
air of their own accord. Lesser Horseshoes and Whiskered Bats 
also moved from positions in which I had observed them, and I 
saw a Long-eared Bat, Plecotus auritus Geoftr., on the wing in 
the evening in one cave. 

On December 29th I carefully noted the position of two Greater 
Horseshoes in the main tunnel of the Long Hole, one of which 
was not more than ten yards from the entrance, but was careful 
not to touch them nor to bring my light neartothem, Twodays 
later both had moved. On the same date (Dec. 29th) I found a 
colony of about forty hanging close together—only a few were 
actually touching any of the others—in the terminal chamber. 
Five of these I took without touching any of the others. When 
pushed from their footholds they fluttered to the ground and lay 
there, cold and lethargic, but squeaking feebly and extending their 
legs backwards, apparently in search of some foothold. The 
remainder drew themselves up by bending their legs but did not 
unfold their wings. Very little dung lay beneath this colony, and 
the bats which I took, though they were soon awake and lively, 


did not defecate in the box; one taken near the entrance of the 
cave did so at once. 

On the 29th I went into the cave in the afternoon, but on the 
31st, when I found that the two bats whose positions I had noted 
had moved, I entered about 6 p.m. A few yards from the 
entrance, in a fissure in the roof which may lead to chambers and 
passages as yet unknown, a thick cluster of wide-awake Greater 
Horseshoes was suspended from a spur of rock. On the floor of 
the cave beneath this fissure I had, on the 29th, noted a large 
pile of dung, but certainly, during the daytime, no bats were 
visible in the fissure. The bats, which numbered at least thirty 
and possibly as many as fifty, were clinging together like a swarm 
of bees; their heads, ears, and facial ornaments were in constant 
nervous movement and the wings hung loosely by their sides; 
they had apparently emerged from some crack or hole in the 
fissure. When I turned the light of a powerful lamp upon the 
bunch, individuals at once detached themselves, dropped on 
outstretched wings, and flew further into the cave. On this 
occasion, so far as I could tell, all flew into the depths of the cave 
and not towards its mouth. J struck two down with my stick as 
they flew and could have killed many more; within afew minutes 
the fissure was empty, ail having apparently entered the cave. 
As I proceeded up the tunnel I found many bats clinging 
to the walls and roof, their wings hanging loosely, their heads, 
ears, and nose-ornaments twitching, as with raised heads and 
necks and hollowed backs they turned and twisted in all 
directions without moving the position of their feet. Whether 
these wide-awake bats were or were not able to see me I am 
not prepared to say, but they were keenly alive to my 
presence and moved, flying freely, when I approached them. 
Other bats passed me, flying up and down the tunnel, but it is 
probable that there is either a second exit or further chambers, 
for I certainly did not come across all the bats which had left the 
cluster near the entrance. In the terminal chamber, where on 
the 29th I had seen about forty bats, only eight remained, and 
these again were awake; while I was watching them two took wing. 

On January Ist, 1907, I went in in the daytime, and not only 
found that the six bats left on the evening before in the terminal 
chamber had disappeared but could not see a single bat hanging 
in the cave. On January 3rd I entered about 6.15 P.m.; no bats 
were visible in the fissure near the entrance where I had seen the 
cluster on December 31st, nor were there any hanging and oniy one 
was on the wing in the tunnel. The chamber at the foot of the 
chimney was again empty, but by climbing over the chock stone 
I reached the upper and larger chamber which I had not previously 
visited. Here I found two small colonies or collections of bats— 
numbering twelve and eight individuals respectively—and a 
single bat in the low passsge which leads out of this chamber. 
These bats were not in profound slumber; some two or three 
individuals were hanging with the wings by their sides and were 

316 MR. T. A. COWARD ON THE -Avor. 95 

in nervous motion. I took the single bat, but did not touch nor 
otherwise disturb the colonies. 

On January 5th, with Mr. Oldham and my usual assistants I 
entered the Long Hole about 5.20 p.m.; bats were then dropping 
out of the fissure near the entrance singly, but there was no cluster 
as there had been on December 31st. About half a dozen came out 
and flew inwards. A few were on the wing in the tunnel and the 
two colonies in the upper chamber were reduced to one company 
of ten individuals. We again left this colony undisturbed. 

In the afternoon, on January 6th, we again visited the upper 
chamber and found that the number of the bats was further 
reduced; only four remained. No bats were flying in the cave 
between 3 and 4 P.M. 

We then returned to the fissure, where I had seen the cluster 
and whence on the previous days we had both seen bats emerge. At 
4.20 p.m. a single Greater Horseshoe dropped out of the hole and 
flew into the cave, and at 4.40 another dropped from the fissure 
and flew out into the open air. Mr. Oldham then posted himself 
at the entrance to the cave, in such a position that he could 
command a view of any bats passing out, while I remaimed below 
the fissure. Between 4.40 and 5.20 p.m. at least nine bats came 
from the fissure or elsewhere and flew out of the cave. A few 
bats passed us in the tunnel; we could hear the whirr of their 
wings distinctly, and before it became too dark to distinguish 
objects at a distance of a few feet—at about 5.30—bats passed. 
and repassed the entrance to the cave, perhaps some of those which 
had emerged intending to re-enter. So long as there was light it 
was easy to see the broad wings of the Greater Horseshoes, as they 
passed out or fluttered past the entrance, silhouetted against 
the sky. 

My visits to the other caves were not so frequently repeated. 
On December 29th I did not observe any Greater Horseshoes in 
Great Oons Cave, but I noted the position of two Lesser Horseshoes 
and a Whiskered Bat without touching them ; the Whiskered Bat 
was clinging to the roof within full reach of daylight. One of the 
Lesser Horseshoes was sleeping lightly; it partially unfolded its 
wings, raised its head, and moved its ears, and then relapsed into 
slumber, refolding its wings. Not one of these three bats, on 
January 5th, was occupying the position I had seen them in on 
December 29th, but one Lesser Horseshoe and a Whiskered Bat 
were in places which were not occupied on the previous visit. 
We found three Greater Horseshoes, two of which were not at the 
spots they occupied on January 5th, when I visited the cave on 
December 29th. 

One large cavern was visited on several occasions; on De- 
cember 31st I could see a number of Greater Horseshoes hanging 
singly in different parts of the cave, but I only took or touched 
one; on the subsequent visits I did not see a single bat. 

About 7 p.m. on December 31st there were at least thirty or 
forty Greater Horseshoes, hanging in two companies some 


distance apart, from the wall of the Echo Cave; the majority of 
these bats were awake and others were on the wing. On 
January 5th the number was smaller; most of these bats were so 
close together that six were dislodged at once by a stone thrown 
from below. The Lesser Horseshoes in this cave may or may not 
have moved in the interval between the visits; they were roughly 
in the same position on both dates. A Long-eared Bat, which, 
when handled and replaced in the same position, appeared to be 
in a deep sleep on the 31st, had gone by the second visit. An 
unidentified bat was on the wing in the daytime just within 
the entrance of Wookey Hole, a cave near Wells. 

Between December 24th, 1906, and January 23rd, 1907, 
Mr. Cummings paid seven visits to the old mines near Barnstaple ; 
he arrived independently at the conclusion that the bats were 
not hibernating and were going outside the caves to feed. He 
has kindly allowed me to add his notes. 

His observations were made at three borings. On December 24th 
he found a single Greater Horseshoe in No. 1, but none in No. 2; 
No. 3, the floor of which was under water, was not entered. On 
December 29th No. 1 was unoccupied, but there were two bats in 
No. 2 twenty-five yards from the entrance. The bats were not 
touched, but when, later in the afternoon, he re-entered this 
boring, he found one on the wing, and the other shortly followed 
the first out of the tunnel. On January 5th there were again two 
bats in No. 2. One occupied a similar position to that when first 
noted, but the other had not returned to the same position. The 
temperature in the open, on the date of this visit, was high, but 
rain was falling steadily. The two bats, after Mr. Cummings had 
taken them out of the tunnel to examine them, and had then 
returned them to their original positions, were roused and driven 
out of the cave; they attempted to re-enter the cave three times, 
but were apparently prevented from doing so by Mr. Cummings’s 
presence at the entrance. Mr. Cummings thought they objected 
to the rain. Later in the afternoon he discovered two bats, which 
showed by their manner of hanging with their wings by their 
sides that they were not in deep sleep; he feels sure they were 
the bats driven out of No. 2. On January 11th No. 1 and No. 2 
were empty, but a small bat entered No. 3 at about 5.20 p.m. and 
shortly afterwards two Greater Horseshoes came out; Mr. Cum- 
mings had, again, not been able to enter No.3. On January 16th 
Nos. | and 2 were again unoccupied ; the temperature in the open 
was on this and the previous visit 48° F.; within the caves 52° 
and 53° F. At 5.30 p.m. two Greater Horseshoes, which came 
from the direction of a deep shaft higher up the slope, passed the 
entrance of No. 38. On January 19th, when the temperature in 
the open had fallen to 40° and a cold east wind was blowing, no 
bats were seen at the mouth of No. 3 between 5.10 and 5.45 p.m., 
neither were any noticed between 5.15 and 5.45 p.m. on 
January 23rd, when the temperature at the mouth of the cave was 
33°, and it had been below freezing-point all day in the open. 

Proc. Zoot. Soc.—1907, No. XXII, 22, 

318 MR. T. A. COWARD ON THE [Apr. 9, 

Mr. Cummings, on this evidence, suggests ‘That the Greater 
Horseshoe, at the end of December and throughout January, 
moves freely in the caves, is easily disturbed, and usually hangs at 
the further end of the boring. That it goes abroad regularly in 
mild weather, but hibernates more or less deeply in frost, and 
remains in the caves during rain or high wind.” 

It has been often asserted that in the Greater Horseshoe and 
other gregarious bats the sexes hibernate apart; though colonies 
consisting of bats of one sex only have been found occasionally, 
both sexes may be found in the same company or cluster. I have 
found male and female Lesser Horseshoes in the same colony at 
Cefn, Denbigh, and of the five bats which I took from the forty 
in the Long Hole on December 29th one was a female and four 
males. The two bats which I killed as they left the cluster at the 
mouth of this cave on December 31st were males; four taken from 
the upper chamber of the Long Hole on January 6th, the remnant 
of the company of twenty, were maies; six dislodged by a stone 
from a congregation on the wall of the Echo Cave were all males; 
and five Pipistrelles, Pipistrellus pipistrellus (Schreber), turned 
out of a crack in the cliff-wall, outside the entrance to the Long 
Hole, were also males. The single bats found in different places 
were of either sex, but the majority were males; their ages 
varied and there were many which, judging by their grey pelage, 
were born in 1906. I may, of course, have failed to discover the 
colonies of females, or some of the bats which we did not touch in 
the colonies may have been of this sex; there is, however, 
another possible explanation, an inequality in the numbers of the 
sexes, the males predominating. 

The Greater Horseshoe feeds in winter, apparently both within 
and outside the caves. The floors of all the caves show evidence 
of the recent movements of bats ; in addition to the piles of dung, 
which were mostly old and dry, fresh excrement was scattered, as 
if dropped by the bats either when flying or hanging from the 
roof. Fragments of the prey, in places in small heaps, were also 
littered about the floors of the caves. 

The stomachs of two Greater Horseshoes, killed immediately 
after capture, contained no food, but that of a third, killed as it 
flew from the bunch at the entrance to the Long Hole, contained 
a little and there was feecal matter in the intestine. 

Large beetles and moths are apparently the chief food of the 
Greater Horseshoe. I found elytra of MJelolontha vulgaris Fabr., 
and the heads and elytra of Geotrupes spiniger Marsh, and perhaps 
G. stercorarius Linn. ; these remains were in most of the heaps of 
dung and were also scattered in the caves; in the Great Oons I 
found them more than 100 or 150 yards from the entrance. Both 
species of Geotrupes are occasionally abroad in mild weather in 
winter : we found one G. spiniger on horse-dung in the road near 
the caves; Mr. Cummings finds Geotrwpes in small numbers 
throughout the winter at Barnstaple. In the Cheddar caves were 
the elytra of the flightless beetle, Vebria brevicollis Fabr., and at 


Barnstaple Mr. Cumimings found fragments of a flightless Ptero- 
stichus (possibly, he thinks, P. niger Schall.). The remains of the 
large cave-spider, Weta menardi Latr.—fragments of the cephalo- 
thorax with the legs attached in some cases—were present in the 
Cheddar caves, and Mr. Cummings found remains of a spider at 
Barnstaple; at Cefn, I found a portion of the leg of a Meta in 
the dung of the Lesser Horseshoe (1). These flightless beetles 
and spiders must have been picked up by the bats from the ground 
or taken from the walls. It is not certain that the bats always 
secure their prey when they are flying, and as the Horseshoes are 
exceedingly agile on the wing I feel sure that they could, by 
hovering, pick food from the ground or from a vertical surface, 
such as the wall of a cave, without alighting ; furthermore, in 
captivity they show a marked tendency to fly low, and will fre- 
quently alight with outspread wings upon a flat surface, springing 
therefrom again with ease. 

The wings of Scotosia dubitata Linn., a moth which hibernates 
freely m limestone-caves and was abundant at Cheddar, were 
scattered here and there, as if eaten recently, and also a few wings 
of Gonoptera libatrix Linn., another species which hibernates, 
were present. These moths might, of course, have been captured 
when they were on the wing in summer, but the remains appeared 
fresher than those of summer-flying moths which do not occupy 
the caves in winter (for example, Zriphena) which I found. 
Mr. Cummings found the remains of both these moths at Barn- 
staple, and also fragments of two large flies, possibly Hristalis 
tenax and some smaller dipteron. At Cheddar there were many 
small dipterous insects resting on the walls of the caves; these are 
soon aroused to activity by the presence of a light. Myr. Robert 
Newstead found, im dung which he kindly examined for me, 
remains of certain J/uscide, which, as he points out, are all diurnal 
insects. We do not, however, know at what hours the bats are 
on the wing in summer, but even if the times when the bats 
emerge and the insects retire in the evening, or vice versd in the 
morning, do not overlap, the bat which can capture a flightless 
beetle or spider could take a fly which was at rest. 

In Goatchurch Cavern, at a great distance from and below the 
entrance, I found a dead Staphylinid beetle of the genus Quedius ; 
it was damaged by the teeth of a bat. 

The prey of the Greater Horseshoe may be captured on the 
wing, but that it is not, as a rule, devoured whilst the bat is flying, 
seems to be proved by the behaviour of bats in captivity even 
more than by the presence of the fragments of prey in the caves. 
When secured by a snap of the bat’s jaws the insect is conveyed 
to some resting-place and there consumed. ‘Two captive Greater 
Horseshoes, one of which survived for 14 days and the other 
for 35 days, were fed almost entirely upon Greotrupes typheus 
Linn., the only beetle I could obtain in sufficient numbers in 
winter. Altogether over 120 beetles were devoured, and in every 

case the behaviour of the bats was practically the same. I usually 

320 MR. T. A. COWARD ON THE [Apr. 9, 

held the bat in my hand until it had snatched the beetle and 
then released it; but at times the bat, when suspended from 
some foothold, would take a beetle which was offered to it. The 
released bat, holding the beetle securely, even if only by one leg, 
invariably flew to some favourite foothold—usually the picture-rail 
in one of three different spots, one end of the window-sill, or a 
curtain-ring—and there hung until the beetle was devoured. 
When the beetle was dropped in flight, only two or three times 
in over 120 experiments, the bat made no attempt to recover its 
prey. I never heard the sound of champing jaws as the bats were 
flying, but when they were at rest the noise of crushing the hard 
armour of the beetles was plainly audible. 

The interfemoral membrane was never used as a pouch, as it 1s 
in the Vespertilionide (2), but the beetle was invariably pushed 
against the interbrachial membrane, as I observed was the case in 
the Lesser Horseshoe (1). At times it was thrust against the 
belly, sometimes into one and sometimes into the other wing, and 
as a rule one leg was detached from its hold in order to give more 
freedom to the half-outstretched wing on the same side. The 
contortions of the bat were most noticeable when it was hanging 

clear and not against the wall; on many occasions one or other of 
my captives went through the whole operation when suspended 
from my finger. The object of this use of the membrane is un- 
doubtedly to prevent the escape of the struggling beetle as the 
bat relaxes its grip in order to adjust the position of the captive 
in its mouth ; when the beetle was seized by the head, especially 
if it was a male armed with sharp thoracic spines, it was turned 
until the bat had a grip upon the abdomen. As the head is 
thrust into the interbrachial membrane, the wing and leg, on the 
same side, are moved sharply forward, thus, by the resistance of 
the membrane, enabling the bat to secure a firmer grasp of the 
beetle, almost as if the wing was used actually to push the prey 
further into the mouth. The object of pouching in the Vesper- 
tilionide is to prevent the escape of a captive, when the bat is 
flying, until a firm grip is secured (2), but mm the Horseshoes the 
use of the interbrachial membrane appears to be different ; in the 
Vespertilionide the head is rapidly withdrawn from the pouch and 
the prey devoured openly ; in the Ahinolophide the wing is made 
use of repeatedly, so long, in fact, as any large portion of the prey 
projects beyond the jaws of the hat. 

The head, prothorax, first pair of legs, and elytra of a beetle are 
usually dropped by the bat, but occasionally the head or one 
elytron are devoured ; those portions rejected by my captives were 
similar to many of the fragments found in the caves. When a 
bat had eaten two or three beetles its head drooped and it lapsed 
into partial slumber, rousing itself spasmcdically at any sudden 
noise. After a few minutes, however, it would be lively again, 
swinging round on its legs and moving its head and ears at the 
slightest sound. 

Mr. J. G. Millais states (8) that this species “ does_not devour 


all its food while in flight, but conveys some to its diurnal resting- 
place”: I will go further; it probably conveys all its food to 
some resting-place. Even when I gave my captives mealworms, 
insects which are easily crushed by the powerful jaws of a Greater 
Horseshoe, they were conveyed to some resting-place unless the 
bat refused to leave my hand. Mr. Millais adds a tootnote : “* Lam 
quite ignorant as to the manner in which these bats carry their 
food to their retreats, since the interfemoral pouch seems to be 
incapable of being bent forward.” In no bats, to my knowledge, 
is the pouch used as a receptacle in which to carry food; the prey 
is carried, in the most natural manner, in the mouth, firmly secured 
by the strong teeth. Mr. Millais’s statement (p. 30) that the tail 
is bent forward and the interfemoral membrane occasionally used 
is probably based on some erroneous observation ; it is the inter- 
brachial and not the interfemoral membrane which is used. During 
the whole process of pouching and eating the beetle the posterior 
portion of the tail remains in the recurved position which is so 
characteristic of the species. 

This reflexed tail, or, to be more exact, portion of the tail, is 
constantly in this curious position. In flight the anterior portion 
of the interfemoral membrane is stretched between the slightly 
flexed legs; the end of the tail is upturned; and when the bat 
is scrambling or climbing the tail is held in the same position ; 
when at rest the tail is flat upon the back if the wings are half 
open, or lies partially concealed by the forearms if the animal is 
closely wrapped in its wings. 

In sleep the position of the wing-membranes is practically the 
same as in the Lesser Horseshoe (4), but, asa rule, the bats I found 
were not so completely encircled by the wings as is the case with 
the smaller species; the forearms seldom met along the back, 
whereas in the Lesser Horseshoe one sometimes overlaps the other. 
In some of the sleeping Greater Horseshoes the tail stood out 
from the back at an angle of about 30°, in a position I have never 
observed in the smaller species. Our knowledge of the habits of 
the Rhinolophide is slight, and as yet we cannot find that the bats 
gain any advantage from this recurved tail ; there is, however, the 
possibility that the tail is rudimentary. 

Occasionally, im captivity, as Mr. Millais noticed, the Greater 
Horseshoe will sleep with the wings folded along the sides, but 
this is an unnatural position and is probably accounted for by the 
impaired health of the captive; when in good health the bats 
sleep with the interdigital membranes one above the other, in a 
similar position to that adopted by some, at any rate, of the 
Pteropodide. The ears, slightly bent or folded, are hidden beneath 
the carpal joint. 

The colour of the adult is brown and does not vary in the sexes, 
but the young are grey; all the young which | found were 
naturally much darker than the August young depicted by 
Mr. Thorburn (5), having more nearly acquired the brown pelage 
of maturity. 

322 MR. T. A. COWARD ON THE [Apr. 9, 

We find, to sum up, the following facts :— 

1. The Greater Horseshoe Bat, and apparently in the mild 
climate of Somerset the Lesser Horseshoe also, if the weather be 
open at the end of December and beginning of January, is not in 
a state of hibernation. It moves in the caves, awakening without, 
artificial stimulus, and leaves the caves apparently in search of 

2. To a great extent the same haunts are used in summer and 
winter, seeing that numbers of both species are to be found 
hanging near the entrances to the caves in the winter in situa- 
tions certainly resorted to during the summer. 

3. Both species are susceptible to the presence of a man when 
they are apparently asleep. They show by their movements—by 
bending their legs and raising and slightly swaying their bodies— 
that they are influenced by the disturbance. It appears, from the 
way in which colonies dwindled in size from time to time, that 
the bats, after being thus disturbed, though not actually awakening 
at the time of the visit, retire from the situations they occupied 
to deeper recesses. 

4. Food is conveyed into the caves from without and devoured 
there, the bats hanging whilst they feed. 

5. Certain creatures are captured and eaten in the caves. 

6. Creatures incapable of flight are captured by the bats and 

7. When feeding the Greater Horseshoe, like its congener, 
makes use of the interbrachial membrane and not of the inter- 
femoral pouch. 

The following is the result of Mr. Robert Newstead’s examination 
of the dung and insect remains which I obtamed in the Cheddar 

Wings of the following moths :— 

Scotosia dubitata Linn. | Probably captured when these insects 
Gonoptera libatriz Linn. } — were hibernating in the caves. 
Triphena orbona Fabr. 
Triphena pronuba Linn. 
NXylophasia polyodon Linn. 

Captured and conveyed into the 
caves in summer. 

Dry dung, obtained in the summer haunt where I found no 
bats in winter. 41 pellets minutely examined. 

About 68 per cent. contained the remains of Lepidoptera. 

ee OO - “ us Coleoptera. 

PES ae es y : Diptera. 

5 aaah: 3 s Arachnida (Spiders). 
4 igs) 5 by Hymenoptera. 

* Pagal tos “ A Trichoptera. 

Of the Coleopterous remains at least 44 per cent. were of 
Geotrupes ; in about 12 per cent. there were fragments of dMelo- 


lontha; in one pellet there were portions of the elytra of a 
Geodephagous beetle, possibly of the genus Amara, and in another 
what appeared to be Dytiscus. 

The fragments of Lepidoptera were difficult to identify, but a 
number of abdominal segments, scales, hairs, antennz, eyes, and 
eges were from moths of the family Noetwidee. 

One pellet was almost entirely composed of the remains of 
Diptera. Mr. Newstead remarks, “including portions of the 
wings of a large Muscid with metallic green body, possibly Lucilia 
or allied genus.” ‘This green fly was pr resent in seven pellets; the 
remains of other Diptera were in three or four pellets. 

In one pellet was ‘“‘a portion of the wing of a Hymenopteron 
remarkably like Vespa.” 

One pellet was entirely composed of the remains of a caddis-fly. 
In December and January I found a caddis-fly on the wing at 
Cheddar, and I picked up the remains of one amongst the 
fragments in the Long Hole. 

The spider-remains were of a large species, probably the cave 
Meta; 1 found remains of this spider, certainly dropped by the 
bats, in the caves. 

Mammalian haivs, probably those of a bat, were present in one 

In 100 pellets, taken from below the fissure in the Long Hole, 
whence we watched bats emerge, about 48 per cent. of the remains 
were of Geotrupes, and a smaller percentage of a large brown 
species, possibly Melolontha. This dung was dry; we may 
conclude, not only from the presence of a summer flying beetle 
(presuming that it was Jelolontha) but from the condition of the 
dung and the large size of the heap, that the fissure is occupied 
in summer. 

Fifty pellets taken from a shallow wide-mouthed cave and 
evidently dropped recently, contained about 98 per cent. of metallic 
fragments of the abdomen and basal segments of the legs of 

One hundred recent pellets from the Long Hole were composed 
of about 90 per cent. of remains of G'eotrupes. 

It would thus appear that during the winter months beetles of 
the genus Geotrupes form the chief food of the Greater Horseshoe 
Bat, and that in summer Lepidoptera and Coleoptera are devoured 
in, approximately, equal quantities. 

The following parasites were on Bats which I obtained at 
Cheddar :— 

Nycteribia hermanni Leach. From two examples of Ahinolophus 
hipposiderus. Identified by Mr. Percy H. Grimshaw. 

Eschatocephalus vespertilionis (C. L. Koch). On &hinolophus 
Serrum-equinum, Rhinolophus hipposiderus, and Myotis 
mystacinus. The Ticks, all females, were identified by 
Mr. A. 8S. Hirst. 

324 MR. F. E, BEDDARD ON THE ANATOMY pavprend, 

Spinturnix vespertilionis (Dufour). This small Mite was present 
on the wing-membrane of Rhinolophus ferrwm-equinum, and 
was identified by Professor E. Trouessart, of Paris, to whom 
Mr. Hirst had kindly forwarded my specimens. 


(1) Cowarp, P. Z.8. 1906, pp. 849-855. 

(2) OupHAM, Zoologist, ser. 4, vol. ui. pp. 471-474 (1899). 

(3) Minnats, Mammals of Great Britain and Ireland, vol. i. 
Ds Ale 
: (4) OtpHam, Manchester Memoirs, vol. xlix. no. 9, pp. 6-7 

(5) Miuuats, op. cit. vol. i. plate 1. 

3. Notes upon the Anatomy of a Species of Frog of the 
Genus Megalophrys, with references to other Genera of 
Batrachia. By Frank EH. Brepparp, M.A., F.RS., 
Prosector to the Society. 

[Received March 19, 1907. } 
(Text-figures 92-100.) 

I possess, through the kindness of Dr. Charles Hose of Borneo, 
an example of a Frog from that country, which at first appears 
to be referable to the genus Megalophrys, and which at least 
comes nearest to—if it be not identical with—Ceratophrys 
nasuta of Schlegel. It appears to differ, however, from that 
Species in one very 1mportant point and in one or two features 
of minor importance. The single example was in an excellent 
state of preservation, and I am therefore able to contribute some 
facts to the anatomy of this genus, which, save for many external 
characters and certain osteological features, is apparently quite 
unknown to Zoologists. It is necessary, however, to preface my 
account of its anatomy with an attempt to fix accurately the 
species to which this account refers. The species I. nasuta was 
originally named and figured by Schlegel*. Our subsequent 
knowledge of the external characters of the species is chiefly due 
to Cantor7, Duméril & Bibron +, Giinther §, Boulenger ||, who 
has summed up and added to the previous descriptions, and, latest 
of all, Werner]. Darwin** has figured the heads of Megalophrys 

* Handi. Dierk., Atlas, pl. iv. fig. 72. The leaf-like appendage upon the nose is 
represented as bemg directed forwards, which is not the case with my example. 

+ Catalogue of Reptiles inhabiting the Malayan Peninsula &c. Calcutta, 1847, 
p. 140. 

{£ Erpétologie Générale, vol. viii. p. 456. 

§ The Reptiles of British India, Ray Soc. 1864, p. 413; Ann. Mag. Nat. Hist. 
(4) xi. p. 419. 

|| B. M. Cat. Batrach. Sal. 1882, p. 443. 

{| “Reptilien und Batrachier aus Sumatra,” Zool. Jahrb. (Abth. f. Syst.) xiii. 
p. 498. ** Descent of Man, 1871, i. p. 26, fig. 32. 


nasuta and M. montana, regarding them, in accordance with the 
first and erroneous view of Giinther, as male and female respectively 
of Megalophrys montana. Kdeling* has figured the entire animal 
from above. All of these descriptions, with the exception of 
that of Boulenger, leave a good deal to be desired in point of 
details of importance, doubtless because those details were 
hardly appreciated as important at the time the papers or works 
in question were written. To Mr. Boulenger’s definition, Werner 
has added a more accurate account of the dermal ossifications, 
the mere occurrence of which was known to his predecessors. I 
shall have occasion to refer again to all the above-named writers 
in the following statement of certain differences which the Frog 
studied by me shows as compared with J/. nasuta. 

The length of the Frog which forms the subject of the present 
communication is 135 mm. from snout to vent, a measurement 
which nearly agrees with that of Werner t, whose description is 
not sufficiently comprehensive to allow of a confirmation of his 
identification, The most salient external characters which point 
to my Frog being identical with Megalophrys nasuta are, in 
correlation with other characters, the long palpebral horn-lke 
appendages and the leaf-like appendage upon the snout. The 
tympanum moreover is concealed, and the loreal region 1s ex- 
cavated. Furthermore, the conical warts agree exactly in their 
disposition with the descriptions given by Cantor, Giinther, and 
Werner ; that is to say, there are three, one on the sacrum median, 
and a pair anteriorly in the shoulder-region. There are, more- 
over, three warts upon the head, all of different sizes, arranged 
in a triangle, the largest occupying the apex of the triangle 
which is directed backwards. Ey (eling ¢, however, whose figure 
of the “species” Megalophrys chysii is not very good, represents only 
the sacral wart and nothing of the kind upon the head. Those, 
indeed, seem to have also escaped the attention of others. No 
characters of value are introduced into his two descriptions of 
the species, and it is impossible to be certain of the identity of 
“ Vegalophrys chysti.” She appendages over the eye and on the 
nose are not of course diagnostic of the genus or species; for 
in various Hemiphractide “there are similar appendages, and 
this family is not to be confused with the Pelobatide. Moreover, 
My. Boulenger has recently § removed from the genus Megalophrys 
and assioned to the allied genus Leptobrachium, “Me galophrys fee ||, 
a species with prominent palpebral processes, the existence of 
which doubtless originally influenced Mr. Boulenger in placing 
the species in that genus. It has also the hard skin upon the 
back which is mentioned as being more strongly developed in 

* Nat. Tijdschr. Nederl. Indié, vol. xxvii. 1864, p. 265 & plate. 

+ Giinther also (Ann. Mag. Nat. Hist. ser. 4, xi. p. 419) speaks of the frog as 
5 inches long. Flower, on the other hand (2: ZS. 1899, p. 916), gives 90 mm. as 
extreme length. 

t Ned. Tijdschr. y. de Dierk. vol. ii. 1865, p. 205, & Nat. Tijdschr. Nederl. Indié, 
vol. xxvii. 1864, p. 265 with plate. 

§ Ann. Mus. Genova, vol. vil. p. 750. || Zbe vol. v. p. 512. 


M. nasuta than in M. montana. In the example studied by 
myself, the calcified (?) area formed a continuous plate beginning 
immediately behind the head, but not continuous with the 
hardened skin lying over the head, reaching back rather beyond 
the anterior ends of the ilia. Laterally this area is bounded 
by a raised glandular fold. On the ventral side of this fold the 
skin is still hardened, though not so much as dorsally ; and this 
area is again bounded by a narrow raised area quite similar to 
that just referred to, which separates it from the soft but tuber- 
cular skin of the belly. The inner finger is distinctly longer 
than the second, a character in which this species agrees with 
M. longipes* rather than IM. montana‘. 

It is part of the generic definition of J/egalophrys according 
to Mr. Boulenger that the outer metatarsals are united. In 

Text-fig. 92. 

Palmar surface of hand (upper figure) and foot (lower figure) of 
Megalophrys nasuta. 

this it differs, according to the same author, from e. g. Pelobates, 
where the “ outer metatarsals are separated by web.’ Having 
been able to compare my Frog with Pelobates fuscus, | find that 

* Boulenger, P. Z.S. 1885, p. 850. + Id., Cat. Baty. Sal. p. 442. 


both frogs agree in the separation by web, only that the web 
is more extended towards the tips of the toes in Pelobates*. In 
this, my species apparently agrees with Megalophrys longipes. 
It has, however, a fairly conspicuous inner metatarsal tubercle, 
but nowhere projecting from the surface of the foot as in 
Pelobates. I found no trace of an outer metatarsal tubercle or 
of subarticular tubercles. These are absent in JZ. longipes, but 
described (together with the inner metatarsal tubercle) as “ in- 
distinct” in MW. montana and (by inference) M/. nasuta. 

There are other characters used for systematic purposes in 
which the individual which I have dissected does not agree with 
its presumed congeners. ‘The vomerine teeth, which are very 
few 7 and form only a narrow band over the projecting region 
of each vomer, are situated distinctly to the inside of and not 
behind each choana. And the loreal region is deeply coneave, as it 
is stated to be in Megalophrys montana and (by inference) in 
M. nasuta by Boulenger =, but apparently not in MW, longipes. 
As a part of the generic definition of M/egalophrys, Boulenger uses 
the form of the tongue, which is described as “ subcircular, 
indistinctly nicked and free behind.’ In my example of 
Megalophrys the tongue is very faimtly$ nicked behind. It is 
also nearly circular and free behind. 

Very important also in fixing the systematic position of this 
Frog are two osteological characters which have been largely 
used in defining the genera of Batrachia. The fusion of the 
sacral vertebra with the ensuing coccyx is a rare feature in 
Batrachian osteology. It occurs, however, in Pipa and among 
the Pelobatide. In two genera only, viz., Scaphiopus and Pelobates, 
does Boulenger describe the sacrum and coccyx as confluent. 
They are most unquestionably so in the Frog which I describe 
in the present communication, and this confluence is incidentally 
shown in the drawing (cf. text-fig. 93) on p. 332, illustrating the 
arrangement of certain muscles. This fusion of the sacral 
vertebra with the coccyx is not only an important classificatory 
fact if existing systems are to be respected, but raises another 
fact of greater Importance. It is well known that among those 
genera of Anura in which this fusion occurs, viz. Pipa, Xenopus, 
and Hymenochirus—the entire set of genera constituting the 
Aglossa—and in Pelobates, Scaphiopus, and the present genus 
among the Pelobatidee, and in Bombinator among the Discoglosside, 
the apparently single sacral vertebra is really double in the case 
of Pipa and Pelobates, formed of three vertebrae in Hymenochirus, 
and of only a single vertebra in Scaphiopus and Bombinator. It 
becomes, therefore, a matter of great interest to ascertain what 
is the arrangement that characterises the sacrum of the species 

* Giinther, Cat. Batr. Sal. 1858, p. 136, speaks of the toes of Ceratophryne nasuta 
as “completely tree.” 

+ Giinther, Cat. Batr. Sal. 1858, p. 1386, writes—“ Vemerine teeth none.” 

{ Cat. Baty. Sal. p. 542. 

§ “ Tongue entire behind.”—Giinther. 


which I provi isionally at least identify with Megalophrys nasuta. 
The most obvious test to apply for the solution of this question 
is the exit of the spinal nerves. Now I find that a stout nerve 
issues from the spinal cord just in front of the stout and expanded 
transverse process of the sacral vertebra; and that an equally 
stout nerve issues behind this transverse process in the angle 
between it and the coccyx. There is none between the two, 
Hence the sacrum is composed of but one vertebra as in most 
other Frogs. 

A second character is the proccelous excavation of the vertebral 
centra, A part of the definition of MJegalophrys by Boulenger is 
“ Vertebree opisthocelian.” This character, used by him in the 
tabular * discrimination of the genera. of ‘Pelobatidee, is again 
made use of in the fuller deser iption of the genus Megalophrys *. 

It is, however, in the characters of the sternum that this 
species chiefly difters from other species of Jegalophrys. Among 
the characters used by Boulenger to define the family Pelobatide, 
one is stated as follows, viz.: “The omosternum is constantly 
present, but small and cartilaginous.” That this definition is 
not absolutely true was subsequently shown by Mr. Boulenger 
himself §, who asserted that in Scaphiopus solitarius there was 
“no omosternum.” The shoulder-girdle and sternum of the 
genus Megalophrys have been figured and deseribed by Parker || 
in Megalophr ys montana. The omosternum is minute and the 
sternum is ossified nearly throughout, an inconspicuous cartila- 
ginous xiphisternum only being left at the free extremity of the 
bone. These characters (¢nter alia) are used by Mr. Boulenger 
in his definition of the genus Megalophrys, and therefore 
presumably also apply to WW. nasuta. Whether the sternal 
apparatus of JZ. longipes has been examined I do not know. In 
the Frog upon which I report in the present communication, the 
characters of the sternal apparatus are different from those 
which have been referred to in MJegalophrys montana. Both 
sternum and omosternum are very well developed. 

The omosternum, instead of being a small oval plate of cartilage 
as it is represented to be by Parker in We galophrys montana, has 

form which apart from details is more like that of Rana 
esculenta. It is actually ten millimetres long, and its proportions 
to the rest of the sternum are therefore much more like those 
of Paludicola bibronti figured by the same author . Although, 
as in WW. montana and other Frogs, the procoracoids are bent very 

* Cat. Batr. Sal. 1882, p. 433. 

+ Ibid. Pp. 442 ; t Loe. cit. p. 432. 

§ P.Z.S. 1899. p. 790. 

|| A Monograph on the “Structure and Development of the Shoulder-Girdle,” 
Ray Soc. 1868, p. 78, pl. vii. fig. 8. 

{| It is by no means certain in every case that the species described by Prof. 
W. K. Parker in his Monograph of the Shoulder-Girdle have been correctly 
identified. I find for example that the sternum of Hyla caerulea (= Calamites 
eyanea” of Parker’s nomenclature) is not as Parker has figured it (Joc. c?é. pl. vii. 
fig. 6), but exactly resembles the sternum of “ Acrodytes daudinii” (presumably 
a Hyla) figured by the same author (Zoe. cit. pl. vil. fig. 1). 



much forwards, their articulation with the omosternum in JZ. nasuta 
is not by any means so far forwards as it is figured by Parker 
in WM. montana. his articulation is plain enough in M/. nasuta. 
Above this articulation and for the entire shaft the omosternum 
is clearly calcified. It expands above into a circular cartilaginous 
plate which has a crescentic outline anteriorly and is wider than 
the shaft. The procoracoidal cartilages at and near both points 
of articulation with the omosternum are also calcified. The 
sternum consists of a bony style as in Megalophrys montana; but 
there are differences in detail. In the first place, the ene 1s 
longer in proportion to its width in W. naswta, and it does not 
expand so markedly in width towards its posterior termination 
as is represented in Prof. Parker’s figure of Megalophrys montana. 
In the second place, J/. montana is characterised by a cartilaginous 
xiphisternum which “is but little extended either laterally or 
axially beyond the shaft-bone.” This is not at all the case with 
Megalophrys nasuta, where the xiphisternum is a broad and 
expanded plate, having posteriorly the semicircular cheese-cutter- 
like outline which is so usual among Frogs. On the whole, 
therefore, there are some differences between the two species. 

The sternum of this Frog is in fact particularly large as 
compared, for example, to that of its ally Pelobates fuscus, with 
which I have compared it in this and in some other details of 
structure. The difference of size is, of course, actual in view of 
the much larger dimensions of the Frog which I describe here as 
compared with the rather small Pelobates fuscus. In Pelobates 
the total length of the body in the individual measured was 
47 mm. and the total length of the sternum 16 mm., the sternum 
being therefore roughly one-third of the length of the body. 
The corresponding measurements in the Frog described here 
weve 135 mm. and 60 mm., the proportions therefore nearly 
approaching one-half. The relationship of the sternum to the 
underlying viscera shows corresponding differences in the two 
genera. In Pelobates the sternum hardly extends back beyond 
the heart and pericardium which, however, it fully covers. The 
liver is left largely exposed. In ‘“ Megalophrys nasuta” the 
sternum passes a considerable distance beyond the liver-lobes, 
the heart being beneath almost the commencement of the sternum 

So far as I can gather from the memoirs already quoted which 
deal with the species Megalophrys nasuta, there has been no 
actual description of the sternum in this particular species. But 
since several systematists use the occurrence of a rudimentary 
omosternum as a generic definition, the matter must have been 
looked into by them, or by some of them. So far as present 
views upon the classification of the Anura go, it is clear that I 
should be hardly wrong in instituting not merely a new species 
but a new genus for this Fr rog, on account of its diver gent sternal 
characters,as compared with those that have already been described 
in the genus Megalophrys. In contradiction, however, to this 


conclusion is the extraordinary similarity in detail-—I particularly 
recall the three warty tubercles upon the back and the ‘“nose- 
leaf”’—between this species and J/. nasuta; but coupled with 
these are also, as it would appear, difterences in features of similar 
ov nearly similar value. Thus the species described here differs 
from J. nasuta, in that the vomerine teeth are within a line 
joining the choanee, and that the inner finger is distinctly longer 
than the second. Again, however, I infer these differences, 
being unable to find any definite description of the characters in 
Megalophrys nasuta. 

The above statement concerning those characters used in the 
discrimination of genera and species among the Anura seems to 
me (unless very grave errors have crept into existing descriptions) 
to show that quite possibly two species have been confused under 
the name of JJegalophrys nasuta, which may even be legitimately 
veferred to different genera. Ido not propose, however, for the 
present to give a nametothe species the anatomy of which I deal with 
in the following pages, in case systematists have really had before 
them, and described, a Frog which is identical with it. But even 
in this case the foregoing description of external characters is not 
without use; for some of these characters have been undoubtedly 
overlooked, or imperfectly described, as in the case of the sternum. 

The above account of the systematic characters of the Frog 
which, on a superficial examination, would be referred to Mega- 
lophrys nasuta as described by Cantor, Gunther, Schlegel, and 
Werner, and, I presume, Boulenger*, may be conveniently sum- 
marised for future reference. The following is a restatement of 
the characters of Megalophrys nasuta based upon the single female 
example of a Frog from Borneo, which, from a survey of the 
external characters as described by several zoologists, would be 
referred to that species ;— 

Length about 5 inches. Head broad and depressed, with loreal 
region excavated. Tympanum invisible. Long palpebral process 
over each eye and an upwardly directed leaf-shaped process on 
the snout. Vomerine teeth few, between choane. Tongue sub- 
circular, very faintly nicked, free behind. Teeth only on upper 
jaw. Three inequisized warts forming a triangle upon posterior 
region of head. Skin of back induraced, like that of head, toa 
joint behind front ends of iia. This region separated by a narrow 
elandular fold from lateral region, also, but more slightly, 
indurated, which is again separated by a similar fold from the 
warty ventral surface. Three prominent tubercles upon back, one 

* Myr. Boulenger’s recent description (Ann. Mus. Genov. ser. 2, iv. p. 512) of a 
Frog, referred to the genus Megalophrys, and to a new species of the same, viz., 
M. fee, which was later (ibid. vil. p. 750) removed by him to the genus Lepto- 
hrachium (with which he also fused the genus Xenophrys), seems to show that 
Mr. Boulenger had not before him at the time when he wrote his ‘ Catalogue of the 
Batrachia Salientia’ examples of the Frog described by myself in the present 
communication. For he has mentioned in describing “ Megalophrys” fee the 
fact that the vertebree are proccelous and that they are opisthoceelous in Meyalophrys 


on each shoulder and one on sacral region. Vertebre proccelous ; 
sacral vertebra fused with coccyx, its transverse processes greatly 
expanded. Noribs. Omosternum not rudimentary, with calcified 
style; sternum a bony style, with expanded cartilaginous xipki- 
sternum ; shoulder-girdle arciferous. Fingers free; inner finger 
longer than second. ‘Toes half-webbed ; outer toe separated from 
next by web. Toes not dilated at tips; no articular tubercles ; 
inner metatarsal tubercle extensive but not separated anteriorly 
from the surface of the foot ; no outer metatarsal tubercle. 

Hab. Borneo. 

It will be observed that in the above brief diagnosis I have not 
attempted to differentiate between generic and specific characters 
or indeed family characters. That this Frog belongs to the 
Pelobatidee is quite plain. That it cannot be referred to any 
known genus—if the present definitions of the same are retained— 
is alsoobvious*. I propose, however, for the present to defer the 
question of generic distinctness until the Pelobatide are better 
known anatomically 7. 

Whether this Frog is a new species or identical with Megalophrys 
nasuta | must, for reasons already stated, leave uncertain. 

§ Muscles of the Back. 

In the various drawings (text-figs. 93-96) submitted herewith in 
illustration of the diaphragm of different species of Batrachia, the 
principal muscles of the back are also shown, and it will be at 
once seen that these differ considerably in the different forms. 
Only in Megalophrys is it impossible to see these different muscles 
(text-fig. 98), for the lengthy transverse muscle of the cesophagus 
and lung almost completely covers them over. When, however, 
this muscle is divided along its greater length, 7. e. across the 
direction of its fibres and the two flaps reflected, the muscles now in 
question are fully displayed as shown in text-fig. 93. The same 
muscles are present which are known to exist in Rana esculenta, 
with the possible addition of a muscle not found either in 
h. esculenta, R. guppy (see p. 333), or 2. tigrina, of which species 
I have examined the last two. The Jntertransversarii muscles 
are well developed and commence from the anterior margin 
of the greatly expanded transverse process of the sacral vertebre. 
They are so well developed that they almost conceal the anteriorly 
lying transverse processes of the vertebrze, each separate band of 
muscle joining successive transverse processes being attached near 
to the ventral median line of the same. The Jlio-lumbaris muscle 

* (Footnote added May 8th.) After the reading of this paper, Mr. Boulenger 
kindly drew my attention to the fact that there occurs in Pelobates cultripes 
a variation In connexion with the fusion or non-fusion of the sacral vertebra 
with the coceyx (‘The Tailless Batrachia of Europe,’ Ray Soc. 1897, part i. p. 208). 
It does not, however, follow that if a character is variable in one species it is not 
a “ good ” character in another. 

+ Schlegel’s name Ceratophryne cannot be resuscitated ; for he first applied it to 
C. dorsata, an American frog which is, I presume, Ceratophrys dorsata. 


is comparatively unimportant, when those muscles in some other 
Frogs to be shortly described are considered. As shown in text- 
fig. 93 its branches to the transverse processes are limited to the 
outer edges of these. The enormously expanded sacral vertebree 
which overlie the iha give off anter lorly the strongest slip of 
muscle which Irefer to the Jlio- lumbaris * complex. <A slender 
slip of muscle represents the origin of the muscle from the ilium. 
This arises from the tip of the ilium as it has been described to 
arise in Rana esculenta; but instead of running forward as in 

Part of dorsal musculature of Megalophrys nasuta. 

es. (Esophageal muscle cut and retlected with oviduct. Coce.I/. Ilio-coccygeal. 
Ii.lumb. Long slip of Mio-lumbaris muscle compared in the text with the 
Musculus proprius pulmonum of Pipa. 8S. Transverse process of sacral 
vertebra, the fusion of which with the coccyx is shown. To the right of 
the long ilio-lumbar slip three short ilic-lumbars are shown overlying the 

that frog, it runs dorsally at right angles to the longitudinal axis 
of the body owing to the overlap of the sacral vertebra to which 
it is partly attached. Jegalophrys differs, as will be seen presently, 
from a number of other Frogs in the presence of a long, almost 
strap-shaped and strong muscle which may perhaps be referred to 
the /lio-lumbaris complex. This arises from the ilium a long way 
back and is continued straight forwards, without any insertions 
of detached slips on the way, to the third vertebra, to the outer 

** This corresponds throughout the following descriptions to the “ Pars lateralis ” 
of Gaupp. 


cartilaginous extremity of the transverse process of which it is 
attached. A flat slip of muscle connects the third vertebra with 
the second, and is exactly in the same straight line with the 
muscle just described. It gives the impression “of being a portion 
of it although it is not actually connected, There is nothing 
exactly com parable to this muscle in any of the genera of Batrachia 
with which I shall presently deal, with the exception of the allied 
Pelobates. Iam inclined, however, to think that it may be the 
homologue of the “musculus pulmonum proprius” of Pipa*, 
which the enormous growth of the transversalis in Megalophrys 
has cut off from communication with the aponeurosis of the lung 
and diverted to the transverse processes of adjacent vertebree. 
But this is at present no more than a suggestion. 

Text-fig. 94. 

Some of the dorsal muscles of Rana guppyi. 
T. Lung. o.d. Oviduct. V. Vertebral centra. Other lettering as in text-tig. 93. 

A comparison between the muscles of Jegalophrys which have 
just been described and those of Rana shows a great number of 
differences. In Megaiophrys the Ilho-lumbar muscle complex is 

* Beddard, P. Z.S. 1895, p. 839, fig. lim. The transversalis sheet to the lung is 
not shown in this figure, ‘ret is in that of Keith (J. Anat. Phys. yxxix. p. 261, 
fig. 13 c). 

Proc. Zoou. Soc.—1907, No. XXITT. 23 


not important as compared with the Intertransversarii, that is if 
the long muscle running from the ilium directly to the third 
vertebra be put out of consideration. In Rana guppyion the other 
hand (see text-fig. 94) the ilio-lumbar muscles are very important 
and occupy a considerable space upon the transverse processes of 
the ribs. The whole muscle appears to te continuous from the top 
of the ilium to the transverse process of the vertebra. But it is, 
though strictly continuous, interrupted by tendinous intersections, 
each one of these corresponding to a transverse process. These 
tendinous intersections are of some width. In Rana tigrina, which 
I have been able to compare, the muscle has the same general 
arrangement; but the tendinous intersections are hardly 
noticeable. This difference is very possibly merely one of those 
differences which may be fairly put down to size, In the larger 
Rana guppyi the complication of the muscle is greater than in its 
smaller ally. 

Text-fig. 95. 


GQ See il 

A dissection of Pelobates fuscus, to show large esophageal sheet of the 
transversalis. Lettering as in text-figs. 98, 94. 

Pelobaies shows a distinct likeness to Megalophrys in the dis- 
position of the muscles now under consideration, as might be 


expected from other points of likeness between the two genera, 
which are referred by some systematists to the same family of 
Aveifera, The striking feature about the muscles of this region 
of the body in Pelobates, is their very massive development as 
compared more particularly with Bufo and Ceratophrys, which I 
shall deal with later on in this communication. When the viscera 
are pushed to one side to render visible these various muscles, the 
ilium and the transverse processes of the vertebree are nearly 
completely invisible. The latter indeed are entirely so; but a 
portion of the ilium is exposed where the massive ilio-coccygeal 
muscle diverges from it. This is by no means the case with other 
genera, as the series of figures (text- figs. 93-96) illustrating these 
facts shows. In this feature again Megalophrys comes nearer to 
Pelobates. But in AMegalophrys the expanded sacral vertebre are 
left uncovered; they are quite covered in Pelodates. 

The strong J/o-coceygeal muscle covers nearly the whole of the 
ilium, arising up to its very tip. To the imside, ¢. e. towards the 
centre of the vertebral column, it conceals the commencement of 
the intertransversarius. The latter muscle is also very strong 
and thick. Instead of lying between adjacent transverse pro- 
cesses, It is not on their plane at all, but entirely covers them and 
moreover shows no visible tendinous inscriptions, Quite anteriorly 
at the second or third vertebra, this muscle finally meets the 
equivalent of the peculiar muscle described above in Jegalophrys. 
That muscle, as in AMegalophrys, avises in Pelobates a long way 
down the ium; it is strap-shaped, but relatively stouter than in 
Megalophrys. In front of the iltum it had no attachment to— 
sent down no slips to—the transverse processes of any vertebre 
save the anterior vertebra to which it is ultimately attached. I 
could detect no ilio-lumbaris other than this muscle, even on 
removing it from its close contact with the intertransversarius ; 
nor indeed is there much room for one. 

So far as the material upon which Tam able to report here enables 
me to say, Ceratophrys stands at one end of a series commencing 
with Pelobetes, and of which Rane is an intermediate member in 
so far as concerns the muscles of the back which are here dealt 
with. Im Ceratephrys the muscular development of this region 
of the body is but feeble, relatively speaking. The transverse 
processes of the vertebree and the larger part of the ilium are 
quite uncovered by muscles, which are extremely shrunken as 
compared with those of Megalophrys and Pelobates and even Rana. 
The same muscles precisely can be recognised and with but little 
modification. The Iho-coceygeal is much reduced and arises only 
from the internal edge of the ilium, commencing some way below 
the tip of that bone and not extending on to the ventral side. 
The Intertransversarii are quite separated by the successive trans- 
verse processes. There is no continuous band of musculature as 
in Pelobates and, though to a less extent, in some of the other types 
described here. The Hio-lumbaris is peculiar as compared with 
that of other genera: it arises as a thick band of muscle from 



near to the tip of the ilium. It forms a continuous mass without 
tendinous intersections, giving off, however, slips to three trans- 
verse processes only, upon the most anterior of which the muscle 

Text-fig. 96. 

Some of the dorsal muscles of Ceratophrys ornata. 

Lettering as in text-figs. 93 & 95. 

In Ceratophrys the vertebre thus suppled by branches of the 
ilio-lumbaris muscles are Nos. 5, 6, 7; but a few fibres are seen 
to run on to the transverse process of the fourth vertebra. 

Bufo closely resembles Ceratophrys in all these features. The 
ilia and the transverse processes of the vertebree are but little 
covered with muscle—at any rate in Lufo marinus and B. agua, 
the two species studied by myself. The ilio-coccygeal is, in pre- 
cisely the same way, a much reduced muscle, commencing its 
origin some way back upon the ilium and not extending upon the 
ventral surface of that bone. The ilio-lumbaris is if anything a 
rather slighter muscle than that of Ceratophrys. Otherwise it is 
exactly similar in its appearance and relations to Bufo martinus 


(I am not quite certain as to B. agua); its fibres are inserted on to 
the transverse processes of vertebree 4 to 7 inclusive. It is from 
the fourth vertebra in these species that the esophageal and lung 
muscles arise (see below, p. 346). In ana also it is from the 
fourth vertebra. 

The resemblance between Bufo and Ceratophrys in the muscles 
now under consideration, particularly in their feeble development 
as compared with the corresponding musclesin some other genera, 
might be put down to similarity of habit. For Ceratophrys 1s 
emphatically a ‘toad’ in its terrestrial habits and lethargic life. 
1 find, however, that there is a close likeness in respect of these 
various structures between Ceratophrys and Leptodactylus which 
is important in view of their inclusion in the same family Cysti 
enathide by some systematists. JT have examined two individuals 
of Leptodactylus hevadactylus, which agree entirely in all the 
structural features to be referred to immediately. A considerable 
portion of each ilium is left perfectly free of muscle. As in 
Ceratophrys, but not in Bufo, the anterior extremities of the ila 
do not reach as far as the transverse process of the eighth vertebra ; 
they extend beyondthem in Lufo. The ilio-coccygeal commences 
some little way down the ilium and gives off slips to the transverse 
processes of vertebre 4 to 8 inclusive, and the origin of the lung- 
muscle is confined to the fourth vertebra. 

There is not, in fact, much difference in these particulars 
between Leptodactylus and Rana. For though in the large Rana 
guppy? the ilium is well covered by the ilio-coccygeal muscle, this 
is not the case with Rana tigrina. And the well-marked tendi- 
nous intersections of the muscle in the larger Rana are not seen 
in the smaller species, which thus resembles Leptodactylus. In 
Ceratophrys the modifications as to these muscles have gone still 
further, and it is perfectly clear from the figures (text-figs. 94, 96) 
that that genus shows marked, though in reality superficial 
differences from other Frogs, which may perhaps, in view of its 
likeness to Bufo, be looked upon as adaptive. It 1s, however, 
perfectly clear that Megalophrys and Pelobates, though differing 
from each other in various details, are on the whole much more 
different from the other genera treated of here than are any of 
those genera among themseives. 

S$ Muscles of the Ventral Surface. 

In the sternal region of MJegalophrys nasuta the following 
muscles are displayed when the skin is removed :—(1) Pars 
abdominalis of Pectoral ; (2) Pars sternalis of Pectoral ; (3) Pars 
epicoracoidalis of Pectoral, (4) Pars episternalis of Deltoid ; and 
(5) Pars scapularis of Deltoid. As contrasted with Rana, the 
most salient difference shown in this dissection is the very large size 
of the episternal head of the deltoid, which is as large as, and of 
course longer than the scapular portion, and together with the 
epicoracoidal head of the pectoral completely hides from view the 


coraco-radialis. The latter is fully exposed in Rena owing to the 
slenderness of the slip, which represents in that Frog the very 
large pars episternalis deltoidei of Megalophrys. 

The Coraco-brachialis brevis seems te be exactly as in Rana. 
The coraco-brachialis longus is a double muscle, exposed only by 
cutting the pars sternalis of the pectoral. 

I could not find the pectoro-cutaneous muscles which are so 
conspicuous in the large Rana guppyt. 

Rectus abdominis.—This muscle arises precisely as in Rana 
from the pelvis, and has here the same conical form, expanding as 
it does rapidly from behind forwards. Between the two Recti 
lies the anterior abdominal vein, which is evident until it dips 
beneath the backwardly prolonged sternum in front. The rectus 
abdominis has only four Jiscriptiones tendinee. Three of these 
lie behind the posterior end of the sternum. The fourth les 
beneath the expanded xiphisternum, covered by it, that is to say, 
when the frog is viewed from the ventral aspect. In Rana 
temporaria there are five of these transverse septa, between the 
several sections of the muscle. Anteriorly to the first inscriptio 
tendinea the main mass of the muscle passes without a break, as 
in Rana, mto the sterno-hyoid. A portion of the rectus muscle, 
however, just anterior to the origin of the pectoralis abdominalis 
and largely covered by the obliquus, is separable by a distinct gap 
and hes at the outside of the main mass of the rectus. This 
muscular slp ends in a strong tendon which is attached to the 
rhomboid swelling of the sternum and seems even to reach the 
coracoid beyond it. 

The Sterno-hyoideus is rather more complicated than in Rana. 
It is first of all found as the forward extension of the rectus 
abdominis. From the expanded xiphisternum—from its anterior 
edge—a flat band of fibresruns forward which dip under the tendon 
of the sternal insertion of the rectus and join the main mass of 
the sterno-hyoid. There is a third origin of this muscle from the 
concealed (dorsal) surface of the sternum. The fibres of this 
muscle arise from further back along the sternum almost from 
the very extremity of the expanded xiphisternum. The more 
anteriorly arising fibres form a separate muscle, distinguishable 
even by a lighter colour, which runs along the inside of the main 
body of the sterno-hyoid. The insertion upon the body of the 
hyoid is more extensive than in either Rana or Pelodytes. The 
insertions of the two muscles are in contact in the middle line of 
the cartilage and they extend further anteriorly. 

§ Hyoid and its Musculature. 

The Hyoid and its musculature ave in some ways peculiar in 
Megalophrys nasuta as compared with those of other Batrachia. 
On opening the body-cavity, a strong pillar of muscle is seen 
anteriorly resting on the pharynx on either side. This is the 
ceratohyal with the enveloping hyvglossal muscle. In Aana the 


ceratohyal is a-continuation on the same plane and in the same 
straight line of the rest of the hyoid. In Megalophrys this part 
of the hyoid arch is bent down at nearly a right angle with the 
rest of the hyoid. The close connection of the bone and its 
ensheathing muscle with the pharynx led me to assume at first 
an actual anatomical connection between the two. A careful 
examination, however, shows that there is no such connection, 
and that the hyoid with the muscle can be raised up from its 
position which is in actual contact with the pharynx. It is even. 
possible that pressure may be exerted upon the pharynx by the 
muscle. In the second place, this hyoglossal muscle is very much 
thicker and altogether stouter than the same muscle in the very 
much larger frog Rana guppyt. As in Rana, the two halves of 
the hyoglossal muscle fuse and become one, and this is continued 
forward in close contact and on the median line of the hyoid to 
the tongue. 

The Submentalis has the usual positions and relations. What. 
is remarkable about it in the present species is that it is actually 
(and therefore @ fortiori relatively) larger than the same muscle 
in Rana guppy. 

The Petrohyoidei ave, asin Rana esculenta*, four in number. 
As in that frog, the petrohyoideus of Megalophrys, arising also 
from the body of the byoid, is larger than the three following 
divisions of the muscle which spring from the bony thyrohyals. 
The last of these in J/egalophrys is a large fan-shaped muscle 
which is not so very much smaller than the anterior petrohyoid. 
And, moreover, the three divisions of this posterior petrohyoideus 
muscle are all of fair size, and not merely in contact at their 
origins (which they are not in Rana esculenta), but the middle 
one actually overlaps the other two. It is important to note that 
in Pelodytes punctatus, according to Ridewoody, the fourth 
division of the petrohyoid is absent. 

The third division of the Petrohyoideus posterior arises, it may 
be remarked, from the inner edge of the thyrohyal and not as the 
other two from or near the outer edge of this bone, 7.e., that 
facing towards the attachment of the muscles. In Rana, Gaupp 
describes the muscle in question as springing from the cartilaginous 
epiphysis of the thyrohyal. This is not at all the case with 
Megalophrys, where the epiphysis in question, easily detached, is 
a relatively long and spur-like plate of cartilage directed outwards 
and at right angles to the shaft of the thyrohyal. Tbe anterior 
ot the three posterior petrohyoids does not arise from the bony 
thyrohyal, but from the cartilage at its Junction with the body of 
the hyoid. 

The Omohyoid appears to show no peculiarities as compared with 

The Geniohyoid has the usual two origins, and it is to be noted 
that these are as in Pelodytes according to Ridewood, and not as 

a Gann s edition of Ecker’s ‘ Anatomie des Frosches,’ Hay, p. 139. 
+ P.Z.S. 1897, p. 579. 


Rana according to the same observer; that is to‘say, its origin 
does not extend on to the tough membrane which lies between 
the posterior process of the body of the hyoid and the thyro- 
hyal. Nor, in Megalophrys, as again in Pelodytes but not in 
Rana, is any part of the sternohyoid inserted on to this membrane. 
The reason for this is clear in the case of Megalophrys. For in 
that Frog the very large posterior petrohyoid completely covers 
over this space and would allow of the insertion of no muscle 
‘upon it. 

The Subhyoideus (the posterior portion of the submaxillaris of 
some authors) is rather different in the Frog which forms the 
subject of the present communication from the corresponding 
muscle of Rana. In the latter it is a hyoidean muscle and arises 
in the extreme lateral region of the hyoid cornua. In Megalophrys, 
where these cornua are absent, I traced the muscle to the wall of 
the skull just behind the tympanum. 

The Hyoid cartilage is not like that of either Pelodytes or Pelo- 
bates, both of them allies of the present genus. Nor does it recall 
that of Rana, for I can find no trace of a cartilaginous cornu 
principale or hyoid arch proper ; and as I have already stated, the 
subhyoideus muscle arises from the skull-wall and did not serve to 
guide me to a hyoid bar. 

Xenophrys monticola obviously comes nearer to my species than 
any other species of the hyoid of which I can find a description. 
Of this Frog the late Prof. W. K. Parker wrote* : “ I could find no 
cartilage in the hyoid arch from the Hustachian opening down- 
wards until I reached the hypohyal region.” The outline is very 
sunilar to that of Megalophrys nasuta. There is towards the 
upper end of the diverging body of the hyoid (the hypohyal of 
Parker) a nick in the cartilage on the outer side. This, as I 
think, represents the nick which I figure here in Megalophrys 
nasuta, and to which what appear to be vestiges of the hyoid 
arches or ceratohyals are attached. Behind this in Yenophrys 
monticola, as in Megalophrys nasuta, the body of the hyoid forms 
a bay from which arises in my species the anterior petrohyoideus. 
There is no more trace in Yenophrys than in my species of an 
anterior lateral process of the body of the hyoid, unless indeed 
the hypohyal really represents that, and the true anterior process 
of the body of the hyoid is absent. In both these Frogs, however, 
the posterior lateral process of the hyoid is present, and the thyroid 
bar is well-developed and ossified, but without the spur-like carti- 
Jaginous epiphysis of Megalophrys nasuta. 

The accompanying figure (text-fig. 97) shows the hyoid of Mega- 
lophrys nasuta. ‘The body is fairly stout and shows no traces of 
an anterior lateral process. The anterior, ‘‘ hypohyal,” process on 
each side is a somewhat spoon-shaped piece of cartilage in the 
same straight line with the rest of the body of the hyoid. It is 
not bent inwards as in Pelobates and Pelodytest. Nor could I 

* “The Skull in the Batrachia,” Phil. Trans. 1881, pl. 28, fig. vil. 
+ P.Z.S. 1897, pl. xxxv. figs. 10, 12. 


find any foramen in the cartilage such as occurs in Pelodytes. 
‘There was a slight thinning, however, of limited extent at a point 
just opposite to the divergence of the two anterior prolongations 
of the body of the hyoid, and corresponding therefore to the 
marked foramina shown in the two genera Pelodytes and Pelobates 
by Dr. Ridewood, whose figures are referred to below, and in 
Scaphiopus*. If that be so, it would seem to follow that the tract 
of cartilage lying outside of this is really the remains of the 
hyoidean cornu, and the apparent solidity of the body of the hyoid 
due to the complete and secondary fusion of this hyoidean cornu 
with the body of the hyoid. The weak place in the body of the 
hyoid is, however, so little marked that I hesitate to give to it 
this morphological importance, and, moreover, it is covered by 

Text-fig. 97. 

Hyoid of Megalophrys nasuta. 

ep. Cartilaginous epiphysis. 7. Ligament representing anterior cornua. 

the attachment of the sterno-hyoideus muscle which lies to the 
inside of the foramen in Pelodytes punctatus. More likely to 
correspond to traces of the otherwise missing hyoidean cornu is, I 
think, a stout ligament shown in my figure (text-fig. 97) which is 
attached to a slight indentation near to the anterior end of the 
anterior prolongation of the body of the hyoid. The position, it will 
be observed, is by no means unsuitable to such an interpretation 
of its nature. Nor would thisargument be necessarily at variance 
with the supposition that the thinning on the body of the hyoid 
in this frog is the equivalent of the foramen in Pelodytes and 

* Boulenger, P. Z. 8. 1899, p. 792. 


Pelobates. 1 could find no splint-bone, such as Ridewood has 
figured in Pelodytes as lying upon the posterior edge of the body of 
the hyoid among the fibres of the sterno-hyoid muscle whose in- 
sertion reaches quite to this edge. The body of the hyoid contained 
no ossified tracts such as are sometimes met with in this cartilage 
in other Frogs. The strongly ossified thyrohyals he, as already 
mentioned, at a considerable angle with the hody of the hyoid. 
The cartilaginous epiphysis of the same is not a cap at the free 
end, but is attached to the outer edge and is somewhat curved. 
It suggests a separate but rudimentary upper element of this 
branchial arch—if it really be one, and not a mere process of the 
body of the hyoid. Dr. Ridewood quotes Mr. Boulenger’s opinion 
‘that the hyoidean cornua were disjointed in all those genera 
which he includes in the family Pelobatide.”* My own results do 
not enable me to support fully this assertion. I would rather say 
that among the Pelobatidee the hyoidean cornua are clearly on the 
road to disappearance. Dr. Ridewood has pointed out several 
points of likeness between these structures in the Aglossa and in 
the Pelobatide. I may add that the disappearance of the anterior 
cornua in Megalophrys nasute and Xenophrys monticola is also 
to be observed in Pipa. 

§ Muscles of Shoulder and Arm. 

The muscles arising from or inserted into the scapula appear 
to be the same in Megalophrys as in Rana; for I identified the 
following, viz.:— Dorsalis scapule, Rhomboideus anterior, Kh. pos- 
terior, Levator scapule superior, L. s, inferior, Cucullaris, Serratus 
superior, S. medius, S. inferior, Interscapularis, and Omohyordeus ; 
besides others of which a further account seems to be desirable. 
There are two of these, viz., the deltoid, and a muscle of which I 
find no account in Gaupp’s edition of Hecker, and which I term 
scapulo-humeralis. 1 have already referred to the pars episternalis 
of the deltoid. The only further point to be remarked upon is 
the apparently total absence of a clavicular head, which head I 
found to be very conspicuous in the large Rana guppy. 

The scapulo-humeralis is a small slender muscle running from 
the scapula to the neck of the humerus, between the scapular and 
adjoining humeral heads of the anconeus. I found this muscle in 
Rana guppy. 

§ Muscles of Thigh. 

The thigh-muscles of Megalophrys are in many respects different 
from those of Rana. 

The most salient difference is seen on the inside of the thigh 
when the skin is removed, and concerns the Semitendinosus. ‘This 
muscle is then evident without further dissection, and is not at 
all covered by the two Graciles. The tendon of insertion of these 
two latter muscles in fact, instead of passing over the tendon of 

* Journ. Linn. Soe. vol. xxvi. p. 53. 


the semitendinosus (that is of course when the thigh is inspected 
from the inside), passes below it. The arrangement is thus the 
precise reverse of that which characterises Rana. 

The Semitendinosus is then quite a superficial muscle on the 
inside aspect of the thigh, visible for practically its whole extent 
without dissection. In Mana it is covered not only by the graciles 
but by the adductor. The semitendinosus may be termed a 
two-headed muscle, but there are traces of further subdivision at. 
the origin from the symphysis. ‘The lower (posterior) half of the 
muscle arises by two closely contiguous heads and is to some 
extent overlapped by the gracilis major. It ends at about the 
end of the second third of the femu: in a stout tendon which is 
continuous with the insertion on to the inside of the leg below 
the knee. The anterior half of the muscle is inserted on to this 
tendon soon after it is established. There is no common fleshy 
origin with any part of the adductor mass such as occurs in Rana. 

Unless the anterior head of the semitendinosus is to be 
regarded as its equivalent, there is no Sartorius muscle. It 1s 
important to notice that in the absence of a sartorius and in the 
superficial semitendinosus, J/egalophrys not merely differs from 
Rana but agrees with Pipa*. 

§ The Lungs and the “ Diaphragm.” 

The Zungs of this species are not much less in size than those 
of the huge female Rana guppyi. The suspensory ligaments 
appear, however, to be rather different and morecomplex. I shall 
give some account of them rather asa basis for future comparison, 
since but little seems to be known concerning these and other 
peritoneal folds among the Batrachia. Anteriorly the right lung 
is not in actual contact with the right lobe of the liver ; ‘there is 
a definite and rather broad pulmo- hepatic ligament, such as is 
pain in Rana guppy. This hgament runs back and at the end 
of the liver-lobe becomes continuous with the postcaval vein. 
It is more extensive than as well as rather different from the 
corresponding ligament in Rana guppyi, and, as will be seen 
presently, extends much further along the lung. This pulmo- 
hepatic and afterwards pulmo-caval ligament is attached to the 
edge of the lung mesiad. It does not end posteriorly in a 
crescentic free edge as such ligaments often do. It runs as far 
as the beginning of the kidney and then bends round and 
becomes counibinomiome with a ligament attaching the dorsal surface 
of the lung t